Understanding Diversity and Systematics in Australian Fabaceae Tribe Mirbelieae

Abstract

Australia has a very diverse pea-flowered legume flora with 1715 native and naturalised species currently recognised. Tribe Mirbelieae s.l. includes 44% of Australia’s peas in 24 genera with 756 recognised species. However, several genera within the Pultenaea alliance in tribe Mirbelieae are considered to be non-monophyletic and two main options have been proposed: option one is to merge ca. 18 genera containing ca. 540 species (the largest genus, Pultenaea has nomenclatural priority); and option two is to re-circumscribe some genera and describe new genera as required to form monophyletic groups. At the species level, option one would require 76% of names to be changed; whereas based on available data, option two is likely to require, at most, 8.3% of names to change. Option two therefore provides the least nomenclatural disruption but cannot be implemented without a robust phylogenetic framework to define new generic limits. Here we present novel analyses of available plastid DNA data (trnL-F) which suggest that option two would be feasible once sufficient data are generated to resolve relationships. However, the reticulate evolutionary histories or past rapid speciation suggested for this group may prevent the resolution of all nodes. We propose targeted use of Next-Generation Sequencing technology as the best way to resolve relationships between the key clades in the tribe and present a framework for such a study. An overview of current taxonomy in the tribe is presented, along with the state of taxonomic knowledge and availability of published descriptions for electronic flora treatments. Several new combinations and typifications are published in an appendix.

Full text

diversity
Review
Understanding Diversity and Systematics in Australian
Fabaceae Tribe Mirbelieae
Russell L. Barrett 1, *, James A. R. Clugston 1, Lyn G. Cook 2, Michael D. Crisp 3, Peter C. Jobson 4,
Brendan J. Lepschi 5, Matthew A. M. Renner 1and Peter H. Weston 1


Citation: Barrett, R.L.; Clugston,
J.A.R.; Cook, L.G.; Crisp, M.D.;
Jobson, P.C.; Lepschi, B.J.; Renner,
M.A.M.; Weston, P.H. Understanding
Diversity and Systematics in
Australian Fabaceae Tribe Mirbelieae.
Diversity 2021,13, 391. https://
doi.org/10.3390/d13080391
Academic Editors: Mario A. Pagnotta,
Mohammad Vatanparast and
Ashley Egan
Received: 30 April 2021
Accepted: 16 August 2021
Published: 20 August 2021
Publisher’s Note: MDPI stays neutral
with regard to jurisdictional claims in
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iations.
Copyright: © 2021 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
1National Herbarium of New South Wales, Royal Botanic Gardens and Domain Trust, Mrs Macquaries Road,
Sydney, NSW 2000, Australia; james.clugston@rbgsyd.nsw.gov.au (J.A.R.C.);
Matt.Renner@rbgsyd.nsw.gov.au (M.A.M.R.); Peter.Weston@rbgsyd.nsw.gov.au (P.H.W.)
2
School of Biological Sciences, The University of Queensland, Brisbane, QLD 4072, Australia; l.cook@uq.edu.au
3Research School of Biology, The Australian National University, Acton, ACT 2601, Australia;
Mike.Crisp@anu.edu.au
4Northern Territory Herbarium, Alice Springs, Department of Environment, Parks and Water Security,
P.O. Box 1120, Alice Springs, NT 0870, Australia; Peter.Jobson@nt.gov.au
5Australian National Herbarium, Centre for Australian National Biodiversity Research, P.O. Box 1700,
Canberra, ACT 2601, Australia; Brendan.Lepschi@csiro.au
*Correspondence: russell.barrett@rbgsyd.nsw.gov.au
Abstract:
Australia has a very diverse pea-flowered legume flora with 1715 native and naturalised
species currently recognised. Tribe Mirbelieae s.l. includes 44% of Australia’s peas in 24 genera with
756 recognised species. However, several genera within the Pultenaea alliance in tribe Mirbelieae are
considered to be non-monophyletic and two main options have been proposed: option one is to merge
ca. 18 genera containing ca. 540 species (the largest genus, Pultenaea has nomenclatural priority);
and option two is to re-circumscribe some genera and describe new genera as required to form
monophyletic groups. At the species level, option one would require 76% of names to be changed;
whereas based on available data, option two is likely to require, at most, 8.3% of names to change.
Option two therefore provides the least nomenclatural disruption but cannot be implemented without
a robust phylogenetic framework to define new generic limits. Here we present novel analyses of
available plastid DNA data (trnL-F) which suggest that option two would be feasible once sufficient
data are generated to resolve relationships. However, the reticulate evolutionary histories or past
rapid speciation suggested for this group may prevent the resolution of all nodes. We propose
targeted use of Next-Generation Sequencing technology as the best way to resolve relationships
between the key clades in the tribe and present a framework for such a study. An overview of current
taxonomy in the tribe is presented, along with the state of taxonomic knowledge and availability of
published descriptions for electronic flora treatments. Several new combinations and typifications
are published in an appendix.
Keywords:
phylogeny; diversification; taxonomy; legumes; biogeography; nomenclature; typifica-
tion; new combinations
1. Introduction
Legumes (Family Fabaceae/Leguminosae) are the third largest plant family behind
daisies (Asteraceae) and orchids (Orchidaceae) and the family contains an incredible range
of morphological and ecological diversity [
1
]. Legumes date back to the Maastrichtian or
early Paleocene, diversifying after the Cretaceous–Paleogene boundary mass extinction
event [
2
]. Estimating the phylogeny and establishing a useful classification system is
naturally a challenging task in such a diverse lineage, however significant advances have
been achieved in recent decades, particularly through contributions to the series Advances
in Legume Systematics [
3
–
14
]. Hundreds of publications have addressed the relationships
between particular clades of Fabaceae using phylogenetic data, and a number of important
Diversity 2021,13, 391. https://doi.org/10.3390/d13080391 https://www.mdpi.com/journal/diversity

Diversity 2021,13, 391 2 of 38
consensus publications have drawn these data together [
15
–
28
]. Broad sampling of genomic
sequence data is now resolving many previously intractable nodes, particularly related to
those nodes that appeared early in the evolution of legumes [
29
,
30
], though it appears that
some nodes may be unresolvable regardless [
31
]. Despite all of these efforts, there are still
many unresolved relationships across the family [25].
Australia’s legume flora of over 3000 species is dominated by the iconic genus Acacia
Mill., with ca. 1100 species [
32
]. The next largest taxonomic group in Australian legumes
is the Faboideae tribe Mirbelieae, with 24 currently accepted genera and 756 recognised
species. The tribe is essentially endemic to Australia, with just a single species (Gomphlobium
nitidum Sol. ex Benth.) extending north to Papua New Guinea [
33
]. Species within the
tribe are distributed across the entire continent, occupying habitats from coastal to alpine
zones, and wet forests to arid deserts. They reach their highest diversity in the South-
west Botanical Province, a biodiversity hotspot, with a secondary centre of diversity
in southeastern Australia [
34
]. Two species, Callistachys lanceolata Vent. and Pultenaea
daphnoides J.C. Wendl., have become naturalised in New Zealand. So far, no taxa in
Mirbelieae are known to have naturalised in South Africa, despite climatic similarities and
other Australian legumes becoming invasive species there [35].
Orthia et al. [
34
,
36
] highlighted the lack of phylogenetic resolution and conflicts
between the available data and existing taxonomic classifications in tribe Mirbelieae,
however, little has been published on the topic since. As these papers were published
over 15 years ago, and little progress has been made since, it is considered timely to
review available data and provide a roadmap for the resolution of both phylogenetic and
taxonomic problems in the tribe. We here combine the available trnL-LF data in a novel
analysis as a framework for discussing known issues and proposing a framework for
future work. While most potential nomenclatural changes must await the generation of
more informative molecular data, a few nomenclatural changes are justified by existing
data and these are made in the current paper. The application of several names is also
clarified through typification to reduce the number of names of uncertain application in
public databases.
1.1. Tribal Limits
Tribe Mirbelieae has been recognised as a natural group for four decades [
37
], sharing
with the closely related tribe Bossiaeeae an endemic distribution in Australia and distinctive
yellow and red corolla markings, hence the common name “egg-and-bacon” peas. Earlier
classifications relied on morphology-based analyses to assess the relationships of genera in
the Mirbelieae (Table 1; [
37
–
41
]). Subsequent studies used DNA sequences to estimate the
phylogeny and interpreted the morphology from its fit to the molecular trees. A molecular
phylogenetic framework for understanding taxonomic limits of the tribes has only partially
been developed [
34
,
36
,
42
–
48
]. Both chloroplast DNA sequences (especially trnL-F) and
nuclear ribosomal DNA (mainly ITS) have been used and are often combined for analysis—
except when conflicting (e.g., [
34
,
48
]). Molecular studies have consistently found both
Bossiaeeae and core Mirbelieae to be monophyletic. These groups differ in embryology [
49
].
A clade comprising core Mirbelieae plus Isotropis Benth. shares reduced, 5-nucleate female
gametophytes as a synapomorphy, while Bossiaeeae share normal 8-nucleate megameto-
phytes but with giant antipodals. The other genera of Mirbelieae s.l., including Daviesia
Sm., Gompholobium Sm., Sphaerolobium Sm., Erichsenia Hemsl. and Viminaria Sm., have
giant antipodals. Relationships among the genera of the giant antipodals group (including
Bossiaeeae) have varied among analyses, and also between the genomes [
46
,
48
]. This
group has usually been found to be non-monophyletic with core Mirbelieae nested inside.
However, in the ITS partition, a base-composition bias (non-stationarity) between groups
with the two embryological traits may explain the non-monophyly [
46
]. When stationarity
was partially corrected by using a Logdet model in a neighbour-joining analysis, the giant
antipodals group was found to be monophyletic, though without branch support [
46
]. That
is, the two embryological groups were reciprocally monophyletic. As monophyly of core

Diversity 2021,13, 391 3 of 38
Mirbelieae has been consistently supported, this tribe could be restricted to the 5-nucleate
embryo-sac group. Then, Bossiaeeae could be expanded to include all the genera with giant
antipodals (i.e., Daviesia,Gompholobium, etc). Another possibility would be to combine
Bossiaeeae and Mirbelieae into a single tribe. In this case, the name Bossiaeeae would
have priority over Mirbelieae. However, reclassification of the tribal boundaries of the
egg and bacon peas is premature until phylogenetic relationships of the genera are better
understood (Table 1).
Table 1.
Genera of Mirbelieae s.l. (column 1); previously published suprageneric classifications (columns 2 and 3); embryo-
sac groups (column 4); usage in the present review; columns 5 and 6 indicate whether genera are monophyletic (yes/no/–)
in published phylogenies with separate cpDNA and ITS trees [36,46–48].
Genus Polhill [37]Genus Group
[46]
Embryo-Sac
[46]Current Paper Monophyletic
cpDNA
Monophyletic
ITS
Jacksonia Mirbelieae Mirbelia group 5-nucleate
Core Mirbelieae
yes yes
Leptosema “ “ “ “ yes yes
Latrobea “ “ “ “ yes yes
Chorizema “ “ “ “ no no
Oxylobium “ “ “ “ no no
Mirbelia “ “ “ “ no no
Callistachys ““““––
Aotus “ “ “ “ no no
Otion ““““––
Urodon ““““––
Phyllota “ “ “ “ yes yes
Euchilopsis ““““––
Dillwynia “ “ “ “ no no
Eutaxia “ “ “ “ yes no
Almaleea ““““––
Stonesiella ““““––
Pultenaea “ “ “ “ no no
Podolobium “ “ “ “ no no
Gastrolobium “ “ “ “ yes * yes *
Isotropis “Isotropis “ Mirbelieae s.l. yes yes
Erichsenia “Daviesia group giant
antipodals “––
Viminaria ““““––
Daviesia “ “ “ “ yes yes
Sphaerolobium “ “ “ “ yes yes
Gompholobium “ “ “ “ no no
Bossiaea Bossiaeeae Bossiaeeae “ Bossiaeeae no no
Platylobium “ “ “ “ yes yes
Goodia “ “ “ “ yes yes
Aenictophyton ““““––
Muelleranthus ““““––
Ptychosema ““““––
Paragoodia ““““––
*: Gastrolobium sensu Chandler et al. [
50
]. “ indicates ‘as above’. ‘–’: multiple species in genus but only one sampled (or genus monotypic).
1.2. Genus-Level Questions
Most recognised genera in Mirbelieae were established in the first half of the 19th
Century [
50
–
67
]. Despite most genera being established for so long, generic boundaries
within core Mirbelieae remain contentious because published phylogenies do not resolve a
number of these genera as monophyletic [
34
,
36
,
68
] (Table 1). Orthia et al. [
34
,
36
] suggested
that the evolutionary history of the core Mirbelieae may be complex, with potential for
ancestral polymorphism, hybridisation, and incomplete speciation/incomplete lineage
sorting. Data also suggest a recent radiation of extant taxa and all of these factors are
potentially resulting in a lack of resolution and statistical support for the major clade
relationships in the tribe. A rapid radiation during the tertiary has been suggested for

Diversity 2021,13, 391 4 of 38
many legume groups [
22
]. Interpretation of phylogenetic histories should take these factors
into account. However, little is known of the presence or frequency of polyploidization
or apomixis in the tribe. Likewise, there is insufficient data to infer any consistency of
maternal inheritance of the chloroplast genome among these genera.
Phylogenetic analyses from the Cook and Crisp labs [
69
–
71
] used both nuclear ITS
data and chloroplast data, including but not necessarily restricted to trnL-F. Trees estimated
from the separate genomes were generally congruent and therefore results from analyses
of combined data were used. However, in an analysis of Pultenaea s.l. [
34
], the chloroplast
and nuclear trees were in conflict and the results were presented separately. Species-level
sampling is still insufficient to resolve a number of generic boundaries, and several key
nodes in the backbone of the trnL-F phylogeny remain unsupported. Some additional data
are available for other markers, including ITS, ETS, trnK, matK, psbA-trnH and trnL, but
sampling density is relatively low. However, these results do provide support for many
clades, including Gastrolobium R.Br., Jacksonia R.Br. ex Sm., Mirbelia Sm., core Oxylobium
Andrews, and core Pultenaea, which contain most species in the core tribe. The resolution
of these key clades as monophyletic provides a high degree of confidence that alternate
classifications can be devised to minimise nomenclatural change once data are generated
that can provide sufficient phylogenetic resolution in order to flesh out the backbone of
the phylogeny. The current lack of resolution means that the best outcome for taxonomic
stability (lumping or dividing existing genera) is uncertain.
One proposed solution is to include all genera in the Pultenaea alliance in a greatly
enlarged mega-genus Pultenaea [
34
], but this has been met with little enthusiasm from the
botanical community and general public. However, an alternative solution is not available
currently, and cannot be obtained without significant new molecular data. If additional
data are generated that resolve relationships, then it is likely that Aotus Sm., Eutaxia
R.Br., Podolobium R.Br. and Pultenaea will require significant re-circumscription. Minor
adjustments are likely to be required in Callistachys Vent., Chorizema Labill. (potentially to
include Podolobium s.s.), Oxylobium and Phyllota (DC.) DC. ex Benth. Several new genera
are likely to be required in order to define monophyletic genera within the tribe, however
this option may require less nomenclatural change than the creation of a giant Pultenaea
including ca. 540 species.
1.3. Species-Level Questions
A significant amount of work has been undertaken to resolve the taxonomic issues
within most genera of tribe Mirbelieae. Full taxonomic revisions have been completed for
Almaleea Crisp & P.H. Weston [
72
], Chorizema [
73
], Daviesia [
69
,
74
–
82
], Erichsenia [
83
], Gas-
trolobium R.Br. [
44
,
71
,
84
–
86
], Gompholobium Sm. [
87
–
89
], Jacksonia [
90
], Leptosema Benth. [
91
],
Phyllota [
92
], Pultenaea [
93
–
97
], and Stonesiella Crisp & P.H. Weston [
42
]. Partial revi-
sions have been completed for Aotus [
98
], Eutaxia [
99
,
100
], Latrobea Meisn. [
101
], Mirbelia
Sm. [102,103] and Sphaerolobium [104–110].
Some genera such as Isotropis are being worked on progressively, with species com-
plexes or distinctive entities being named as they are resolved [
111
–
116
]. Revisionary work
is in progress for the complex genus Dillwynia Sm. [
117
,
118
], and four new combinations
are provided in Appendix Aof this paper, building on three new species published by
Jobson and Weston [
119
,
120
]. Regional flora treatments, including tribe Mirbelieae, are
available for many taxa in the genera mentioned above and all or most species of Callis-
tachys Vent., Euchilopsis F. Muell., Oxylobium Andrews, Urodon Turcz. and Viminaria, though
several of these genera still require systematic revision [
111
,
118
,
121
–
138
]. New putative
species are still being identified on a regular basis in Australia, and many of these new
discoveries are localised taxa that warrant conservation listing.
Despite relatively recent revisions [
93
–
97
], morphological assessments of Pultenaea
s.s. have identified many species circumscriptions that remain uncertain (R.L. Barrett,
J.A.R. Clugston & M.A.M. Renner, unpubl. data). There are also unresolved issues of
typification, as not all relevant material was located during these revisions (R.L. Barrett,

Diversity 2021,13, 391 5 of 38
unpubl. data) We estimate that around a third of the 148 species currently included in
Pultenaea should have their taxonomic circumscriptions revised. Many changes differing
from these treatments have already been adopted by the Flora of South Australia [
124
]
and VicFlora [139]. Detailed assessment of the Pultenaea glabra Benth. complex by M.A.M.
Renner et al. (in prep.), has identified six new species. As Pultenaea glabra is already listed
as rare in its broadest circumscription, all of the segregate species are likely to be listed as
threatened taxa within New South Wales. It is likely that many additional taxa that are of
conservation priority should be recognised in Pultenaea.
1.4. Biogeographic Patterns
Tribe Mirbelieae is of significant interest for understanding the evolution of the Aus-
tralian flora due to its distribution across Australia, high species diversity [
68
,
140
], high
levels of local endemism [
71
,
141
–
143
], morphological diversity [
46
], breeding systems [
144
],
and specialised pollination syndromes [
48
,
145
]. There are contrasting centres of endemism
in both southwestern (particularly Daviesia and Gastrolobium) and southeastern Australia
(particularly Bossiaea,Dillwynia and Pultenaea), a pattern not seen in many species-rich
groups in Australia [
40
,
146
,
147
]. These parallel patterns offer unique opportunities for un-
derstanding broader patterns and processes of speciation in the two regions. These strong
biogeographic trends are reflected in phylogenetic reconstructions of the tribe (Figure 1). A
comparison of south-west and south-east diversity patterns in Daviesia and Bossiaea [
70
]
revealed that the geographic overlap of clades was significantly greater for Daviesia in the
south-west than in the south-east but that this was reversed for Bossiaea. Despite this, diver-
sification rates did not differ between the regions in either genus over the last
10 Myr [70]
.
Rather, the interaction of multiple factors likely explains the diversity differences between
the two regions. The smaller south-western geographic ranges of species in both genera
are probably explained by the steeper climatic gradients in that region. Daviesia is far
more species-rich in the south-west than in the south-east, likely because of its longer
evolutionary time there, combined with the greater geographic overlap of clades [70].

Diversity 2021,13, 391 6 of 38
Diversity 2021, 13, x FOR PEER REVIEW 6 of 39
(A)
Figure 1. Cont.

Diversity 2021,13, 391 7 of 38
Diversity 2021, 13, x FOR PEER REVIEW 7 of 39
(B)
Figure 1. Cont.

Diversity 2021,13, 391 8 of 38
Diversity 2021, 13, x FOR PEER REVIEW 8 of 39
(C)
Figure 1. Cont.

Diversity 2021,13, 391 9 of 38
Diversity 2021, 13, x FOR PEER REVIEW 9 of 39
(D)
Figure 1. (A–D) Phylogeny of Fabaceae tribe Mirbelieae inferred from trnL intron and trnL–F spacer (cpDNA). Posterior
output from MrBayes. Numbers above lines are posterior probability.
Figure 1.
(
A
–
D
) Phylogeny of Fabaceae tribe Mirbelieae inferred from trnL intron and trnL–F spacer (cpDNA). Posterior
output from MrBayes. Numbers above lines are posterior probability.

Diversity 2021,13, 391 10 of 38
2. Materials and Methods
Phylogenetic Analyses
A phylogeny of core Mirbelieae has been inferred here using the trnL intron and
trnL–F spacer of cpDNA. Previous studies have only included selected data, so a novel
analysis is justified in order to assess all of the available data for this marker. All available
sequences for tribe Mirbelieae were downloaded from NCBI Genbank (Table 2). As genera
outside core Mirbelieae are relatively distantly related [
46
], and the key questions we wish
to address relate to core Mirbelieae, only six species outside this core were included as
outgroups (Table 2). Randomly selected sequences were checked by using BLAST (http:
//blast.ncbi.nlm.nih.gov/Blast.cgi (accessed on 25 April 2021)) to ensure that they were
not from a contaminated source. A preliminary alignment was performed using MAFFT
ver. 7.450 [
148
,
149
] and minor corrections were made manually with Geneious Prime
(ver. 2021.1.1, see https://www.geneious.com, accessed on 25 April 2021). Phylogenetic
analyses were performed using Bayesian inference implemented in MrBayes (ver. 3.2.6,
see https://github.com/NBISweden/MrBayes/releases/tag/v3.2.6 (accessed on 25 April
2021)) [
147
,
148
]. Daviesia longifolia Benth. was designated as the outgroup for analyses. Four
Markov-Chain Monte Carlo (MCMC) chains were run for 20 million generations, with one
tree sampled every 5000 generations at a temperature of 0.2 with default priors (gamma),
and GTR substitution model, GTR+I+G (identified by the Akaike Information Criterion
corrected for small sample sizes in jModelTest2; https://github.com/ddarriba/jmodeltest2
(accessed on 30 June 2021)). The first 2,000,000 trees recovered were discarded as burn-in
(trees produced before convergence). Stationarity and mixing were assessed using Tracer
(ver. 1.7.1) [150].
Table 2.
Taxa analysed, vouchers and GenBank reference numbers for the trnL intron and trnL–F spacer. Herbarium codes
for voucher location follow Thiers (2008 onwards; http://sweetgum.nybg.org/science/ih/ (accessed on 25 April 2021)).
Taxon Collector Number Herbarium GenBank
Almaleea Crisp & P.H.Weston
A. cambagei (Maiden & E.Betche) Crisp &
P.H.Weston M.D. Crisp 9197 CANB AF113775
Aotus Sm.
A. carinata Meisn. J.A. Chappill 6581 PERTH AY883181
A. cordifolia Benth. J.A. Chappill 6587 PERTH AY883182
A. sp. Esperance (P.G. Wilson 7904) M.D. Crisp 9197 CANB AF518163
A. subglauca Blakey & McKie M.D. Crisp 9047 CANB AF113776
Callistachys Vent.
C. lanceolata Vent. G.T. Chandler 474 CANB AY015072
Chorizema Labill.
C. aciculare (DC.) C.A.Gardner M.D. Crisp 9202 CANB AF518149
C. carinatum (Meisn.) J.M.Taylor & Crisp M.D. Crisp 9237 CANB AF518150
C. dicksonii Graham M.D. Crisp 9166 CANB AY251250
C. genistoides (Meisn.) C.A.Gardner M.D. Crisp 9026 CANB AF518151
C. obtusifolium (Sweet) J.M.Taylor & Crisp M.D. Crisp 9201 CANB AY251251
C. parviflorum Benth. M.D. Crisp 9166 CANB AF518152
C. rhombeum R.Br. M.D. Crisp 9230 CANB AF518153
C. ulotropis J.M.Taylor & Crisp M.D. Crisp 9214 CANB AY251253
C. varium Benth. M.D. Crisp 8528 CANB AF518154
Dillwynia Sm.
D. parvifolia R.Br. ex Sims J.M. Taylor 360 CBG AF113777
D. phylicoides A.Cunn. M.D. Crisp 9049 CANB AF113778
Euchilopsis F.Muell.
E. linearis (Benth.) F.Muell. M.D. Crisp 8535 CANB AF113779
Eutaxia R.Br.

Diversity 2021,13, 391 11 of 38
Table 2. Cont.
Taxon Collector Number Herbarium GenBank
C. racemosum (Meisn.) J.M.Taylor & Crisp M.D. Crisp 9012 CANB AY251252
E. microphylla (R.Br.) C.H.Wright & Dewar M.D. Crisp 8918 CANB AF113780
E. neurocalvx (Turcz.) Chappill & G.R.Hend. M.D. Crisp 8525 CANB AF113789
Gastrolobium R.Br.
G. alternifolium G.Chandler & Crisp M.D. Crisp 8512 CANB AY015088
G. bilobum R.Br. G.T. Chandler 724 CANB AY015073
G. bracteolosum (F.Muell.) G.Chandler & Crisp G.T. Chandler 426 CANB AY015063
G. brownii Meisn. G.T. Chandler 726 CANB AY015074
G. calycinum Benth. G.T. Chandler 544 CANB AY015075
G. celsianum (Lem.) G.Chandler & Crisp M.D. Crisp 9009 CANB AY015064
G. congestum G.Chandler & Crisp G.T. Chandler 404 CANB AY015076
G. coriaceum (Sm.) G.Chandler & Crisp G.T. Chandler 723 CANB AY015089
G. cuneatum Henfr. M.D. Crisp 8937 CANB AY015077
G. ebracteolatum G.Chandler & Crisp M.D. Crisp 8471 CANB AY015102
G. formosum (Kippist ex Lindl.) G.Chandler &
Crisp M.D. Crisp 8933 CANB AY015085
G. grandiflorum F.Muell. G.T. Chandler 598 CANB AY015078
G. hookeri Meisn. M.D. Crisp 8907 CANB AY015090
G. latifolium (R.Br.) G.Chandler & Crisp G.T. Chandler 365 CANB AY015065
G. leakeanum J.Drumm. M.D. Crisp 8481 CANB AY015091
G. luteifolium (Domin) G.Chandler & Crisp M.D. Crisp 9407 CANB AY015092
G. melanopetalum (F.Muell.) G.Chandler & Crisp M.D. Crisp 8470 CANB AY015066
G. minus (Crisp) G.Chandler & Crisp M.D. Crisp 8922 CANB AY015067
G. modestum (Crisp) G.Chandler & Crisp M.D. Crisp 8465 CANB AY015068
G. obovatum Benth. G.T. Chandler 657 CANB AY015093
G. parviflorum (Benth. ex Lindl.) Crisp G.T. Chandler 760 CANB AY015079
G. plicatum Turcz. 1 G.T. Chandler 623 CANB AY015094
G. plicatum Turcz. 2 M.D. Crisp 9014 CANB AF518161
G. praemorsum (Meisn.) G.Chandler & Crisp G.T. Chandler 729 CANB AY015069
G. pulchellum Turcz. M.D. Crisp 8480 CANB AY015095
G. pusillum Crisp & P.H.Weston M.D. Crisp 8921 CANB AY015080
G. pyramidale T.Moore G.T. Chandler 488 CANB AY015096
G. reticulatum (Meisn.) Benth. G.T. Chandler 540 CANB AY015097
G. sericeum (Sm.) G.Chandler & Crisp J.M. Taylor 1959 CBG AY015070
G. spathulatum Benth. ex Lindl. M.D. Crisp 8448 CANB AY015098
G. spinosum Benth. ex Lindl. G.T. Chandler 548 CANB AY015081
G. subcordatum (Benth.) G.Chandler & Crisp M.D. Crisp 8511 CANB AY015071
G. truncatum Benth. M.D. Crisp 8919 CANB AY015082
G. vestitum (Domin) G.Chandler & Crisp M.D. Crisp 8489 CANB AY015099
Jacksonia Sm.
J. alata Benth. M.D. Crisp 8956 CANB AF518146
J. horrida DC. M.D. Crisp 8934 CANB AY015084
J. macrocalyx Meisn. M.D. Crisp 9272 CANB AF518147
Latrobea Meisn.
L. brunonis (Benth.) Meisn. J.A. Chappill 6564 PERTH AY883186
L. genistoides (Meisn.) Meisn. J.A. Chappill 6567 PERTH AY883187
L. hirtella (Turcz.) Benth. M.D. Crisp 8478 CANB AF113781
Leptosema Benth.
L. aphyllum (Hook.) Crisp M.D. Crisp 9019 CANB AF518148
L. daviesioides (Turcz.) Crisp M.D. Crisp 9193 CANB AY883188
Mirbelia Sm.
M. baueri (Benth.) Joy Thomps. M.D. Crisp 9144 CANB AY251254
M. confertiflora Pedley M.D. Crisp 9050 CANB AF518155
M. depressa E.Pritz. M.D. Crisp 9020 CANB AY015086
M. dilatata R.Br. M.D. Crisp 8491 CANB AY015087
M. longifolia C.A.Gardner 1 M.D. Crisp 9263 CANB AY883189

Diversity 2021,13, 391 12 of 38
Table 2. Cont.
Taxon Collector Number Herbarium GenBank
M. longifolia C.A.Gardner 2 M.D. Crisp 9263 CANB AY251255
M. microphylla (Turcz.) Benth. A. Monro 22 CANB AF518156
M. oxylobioides F.Muell. 1 M.D. Crisp 9112 CANB AY251264
M. oxylobioides F.Muell. 2 Yi 14,825 KUN NC_047371
M. oxylobioides F.Muell. 3 Yi 14,825 KUN MN709855
M. pungens A.Cunn. ex G.Don. M.D. Crisp 9138 CANB AY251257
M. ramulosa (Benth.) C.A.Gardner A. Monro 4 CANB AY251258
M. rhagodioides Crisp & J.M.Taylor M.D. Crisp 9259 CANB AY251259
M. rubiifolia (Andrews) G.Don ANBG 8406509 CANB AF518157
M. seorsifolia (F.Muell.) C.A.Gardner H. King 314 CANB AY251260
M. sp. Pedicellosa (A.Monro 25) A. Monro 25 CANB AY251256
M. sp. Ternata (M.D.Crisp & L.G.Cook MDC
9267) M.D. Crisp 9266 CANB AY251262
M. speciosa Sieber ex DC. subsp. speciosa 1 M.D. Crisp 9123 CANB AY251261
M. speciosa Sieber ex DC. subsp. speciosa 2 – 8100876 CBG AF518158
M. viminalis (A.Cunn. ex Benth.) C.A.Gardner D. Morris 547 CANB AY251263
‘Otion’ Crisp & P.H.Weston ined.
‘O. microphyllum’ (Benth.) Crisp & P.H.Weston
ined. M.D. Crisp 8970 CANB AF113782
Oxylobium Andr.
O. arborescens R.Br. 1 G.T. Chandler 616 CANB AY015100
O. arborescens R.Br. 2 M.D. Crisp 9093 CANB AF113783
O. cordifolium Andrews M.D. Crisp 9133 CANB AF518159
O. ellipticum (Vent.) R.Br. 1 M.D. Crisp 9092 CANB AF113784
O. ellipticum (Vent.) R.Br. 2 G.T. Chandler 603 CANB AY015101
O. lineare (Benth.) Benth. M.D. Crisp 8471 CANB AY015102
O. pulteneae DC. M.D. Crisp 9046 CANB AY015103
O. robustum Joy Thomps. I.R. Telford 4294 CBG AY015104
Phyllota (DC.) Benth.
P. phylicoides (Sieber ex DC.) Benth. 1 M.D. Crisp 9048 CANB AF113785
P. phylicoides (Sieber ex DC.) Benth. 2 M.R. Gillings Soil core – JN392738
Podolobium R.Br.
P. aciculiferum F.Muell. G.T. Chandler 606 CANB AF518160
P. aestivum Crisp & P.H.Weston G.T. Chandler 612 CANB AY015105
P. alpestre (F.Muell.) Crisp & P.H.Weston G.T. Chandler 1039 CANB AY015106
P. ilicifolium (Andrews) Crisp & P.H.Weston G.T. Chandler 308 CANB AY015107
P. procumbens (F.Muell.) Crisp & P.H.Weston B. Hadlow 461 CBG AY015108
P. scandens (Sm.) DC. G.T. Chandler 309 CANB AY015109
Pultenaea Sm.
P. adunca Turcz. J.A. Chappill 6544 PERTH AY883190
P. alea de Kok J. Westaway 924 CANB AY883191
P. arida E.Pritz. 1 J.A. Chappill 6272 PERTH AF518162
P. arida E.Pritz. 2 L.A. Orthia 71 CANB AY883192
P. aspalathoides Meisn. L.A. Orthia 50 CANB AY883193
P. baeuerlenii F.Muell. B. Pfeil 263 CANB AY883194
P. barbata C.R.P.Andrews T.R. Lally 1309 CANB AY883195
P. phylicoides (Sieber ex DC.) Benth. 1 M.D. Crisp 9048 CANB AF113785
P. phylicoides (Sieber ex DC.) Benth. 2 M.R. Gillings Soil core – JN392738
Podolobium R.Br.
P. aciculiferum F.Muell. G.T. Chandler 606 CANB AF518160
P. aestivum Crisp & P.H.Weston G.T. Chandler 612 CANB AY015105
P. alpestre (F.Muell.) Crisp & P.H.Weston G.T. Chandler 1039 CANB AY015106
P. ilicifolium (Andrews) Crisp & P.H.Weston G.T. Chandler 308 CANB AY015107
P. procumbens (F.Muell.) Crisp & P.H.Weston B. Hadlow 461 CBG AY015108
P. scandens (Sm.) DC. G.T. Chandler 309 CANB AY015109
Pultenaea Sm.
P. adunca Turcz. J.A. Chappill 6544 PERTH AY883190
P. alea de Kok J. Westaway 924 CANB AY883191

Diversity 2021,13, 391 13 of 38
Table 2. Cont.
Taxon Collector Number Herbarium GenBank
P. arida E.Pritz. 1 J.A. Chappill 6272 PERTH AF518162
P. arida E.Pritz. 2 L.A. Orthia 71 CANB AY883192
P. aspalathoides Meisn. L.A. Orthia 50 CANB AY883193
P. baeuerlenii F.Muell. B. Pfeil 263 CANB AY883194
P. barbata C.R.P.Andrews T.R. Lally 1309 CANB AY883195
P. blakelyi Joy Thomps. R.P.J. de Kok 715 CANB AY883196
P. brachyphylla Turcz. L.A. Orthia 56 CANB AY883197
P. brachytropis Benth. L.A. Orthia 37 CANB AY883198
P. calycina (Turcz.) Benth. subsp. calycina L.A. Orthia 51 CANB AY883199
P. calycina subsp. proxena Orthia & Chappill L.A. Orthia 63 CANB AY883199
P. costata H.B.Will. R.P.J. de Kok 730 CANB AY883200
P. daena Orthia L.A. Orthia 57 CANB AY883201
P. daphnoides J.C.Wendl. E. Gauba 22,264 CBG AF113786
P. densifolia F.Muell. J. Mant 71 CANB AY883202
P. dentata Labill. M.D. Crisp 9053 CANB AY015110
P. divaricata H.B.Will. F.E. Davies 1887 CBG AY883203
P. echinula Sieber ex DC. R.P.J. de Kok 709 CANB AY883204
P. elachista (F.Muell.) Crisp L.A. Orthia 66 CANB AY883205
P. empetrifolia Meisn. 1 L.A. Orthia 54 CANB AY883206
P. empetrifolia Meisn. 2 L.A. Orthia 53 CANB AY883385
P. empetrifolia Meisn. 3 L.A. Orthia 79 CANB AY883386
P. empetrifolia Meisn. 4 L.A. Orthia 80 CANB AY883387
P. ericifolia Benth. 1 L.A. Orthia 88 CANB AY883207
P. ericifolia Benth. 2 L.A. Orthia 39 CANB AY883388
P. ericifolia Benth. 3 L.A. Orthia 41 CANB AY883389
P. ericifolia Benth. 4 L.A. Orthia 46 CANB AY883390
P. ericifolia Benth. 5 L.A. Orthia 48 CANB AY883391
P. ericifolia Benth. 6 M.D. Crisp 8451 CANB AF113788
P. euchila DC. D.A. Taylor 55 CANB AY883208
P. fasciculata Benth. R.P.J. de Kok 860 CANB AY883209
P. flexilis Sm. R.P.J. de Kok 705 CANB AY883210
P. glabra Benth. R.P.J. de Kok cultivated – AY883211
P. graveolens Tate R.P.J. de Kok 778 CANB AY883212
P. gunnii Benth. R.P.J. de Kok 900 CANB AY883213
P. heterochila F.Muell. 1 J.A. Chappill 4279 PERTH AY883231
P. heterochila F.Muell. 2 D.A. Taylor 1569 CANB AY883401
P. hispidula Benth. R.P.J. de Kok 780 CANB AY883214
P. indira Orthia & Crisp subsp. indira 1 L.A. Orthia 59 CANB AY883215
P. indira Orthia & Crisp subsp. indira 2 L.A. Orthia 70 CANB AY883395
P. indira Orthia & Crisp subsp. indira 3 M.D. Crisp 9178 CANB AY883396
P. indira subsp. monstrosita Orthia L.A. Orthia 78 CANB AY883397
P. juniperina Labill. R.P.J. de Kok 713 CANB AY883216
P. kraehenbuehlii P.J.Lang R.J. Bayer SA99010 CANB AY883217
P. largiflorens F.Muell. ex Benth. D.L.Jones 15,757 CANB AY883218
P. laxiflora Benth. A. Monro 64 CANB AY883219
P. luehmannii Maiden R.P.J. de Kok 738 CANB AY883220
P. muelleri Benth. R.P.J. de Kok 779 CANB AY883221
P. myrtoides A.Cunn. D.A. Taylor 7 CANB AY883222
P. ochreata Meisn. L.A. Orthia 42 CANB AY883223
P. parviflora Sieber ex DC. R.P.J. de Kok 793 CANB AY883224
P. patellifolia H.B.Will. R.P.J. de Kok 761 CANB AY883225
P. pauciflora M.B.Scott 1 L.A. Orthia 83 CANB AY883400
P. pauciflora M.B.Scott 2 L.A. Orthia 84 CANB AY883226
P. pauciflora M.B.Scott 3 L.A. Orthia 85 CANB AY883399
P. pedunculata Hook. R.P.J. de Kok 756 CANB AY883227
P. petiolaris A.Cunn. ex Benth. R.P.J. de Kok 903 CANB AY883228
P. pinifolia Meisn. L.A. Orthia 40 CANB AY883229
P. purpurea (Turcz.) Crisp & Orthia L.A. Orthia 60 CANB AY883230

Diversity 2021,13, 391 14 of 38
Table 2. Cont.
Taxon Collector Number Herbarium GenBank
P. radiata H.B.Will. L.A. Orthia 38 CANB AY883232
P. reticulata (Sm.) Benth. L.A. Orthia 47 CANB AY883233
P. rosmarinifolia Lindl. D.A. Taylor 68 CANB AY883234
P. rotundifolia (Turcz.) Benth. L.A. Orthia 61 CANB AY883235
P. scabra R.Br. R.P.J. de Kok 909 CANB AY883236
P. sericea (Benth.) Corrick J. Mant 72 CANB AY883237
P. setulosa Benth. R.P.J. de Kok 716 CANB AY883238
P. skinneri F.Muell. L.A. Orthia 36 CANB AY883239
P. sp. Mt Lesueur (Beard 7827) J.S. Beard 7827 PERTH AY883398
P. spinosa (DC.) H.B.Will. R.P.J. de Kok cultivated – AY883240
P. spinulosa (Turcz.) Benth. L.A. Orthia 68 CANB AY883241
P. stipularis Sm. R.P.J. de Kok 701 CANB AY883242
P. stricta Sims R.P.J. de Kok 729 CANB AY883243
P. strobilifera Meisn. 1 L.A. Orthia 44 CANB AY883392
P. strobilifera Meisn.2 L.A. Orthia 52 CANB AY883393
P. strobilifera Meisn.3 L.A. Orthia 75 CANB AY883394
P. subalpina (F.Muell.) Druce R.P.J. de Kok 721 CANB AY883244
P. subspicata Benth. R.P.J. de Kok 718 CANB AY883245
P. tarik de Kok R.P.J. de Kok 666 CANB AY883246
P. tenuifolia R.Br. & Sims 1 L.A. Orthia 64 CANB AY883247
P. tenuifolia R.Br. & Sims 2 R.P.J. de Kok 803 CANB AY883248
P. trichophylla H.B.Will. ex J.M.Black R.P.J. de Kok 838 CANB AY883249
P. trifida J.M.Black R.P.J. de Kok 819 CANB AY883250
P. trinervis J.M.Black R.P.J. de Kok 801 CANB AY883251
P. tuberculata Pers. R.P.J. de Kok 702 CANB AY883252
P. verruculosa Turcz. L.A. Orthia 45 CANB AY883253
P. vestita R.Br. ex Aiton R.P.J. de Kok 830 CANB AY883254
P. victoriensis Corrick R.P.J. de Kok 762a CANB AY883255
P. villifera Sieber ex. DC. R.P.J. de Kok 828 CANB AY883256
P. viscidula Tate R.P.J. de Kok 833 CANB AY883257
P. vrolandii Maiden R.P.J. de Kok 784 CANB AY883258
P. whiteana S.T.Blake M.D. Crisp 9113 CANB AY883259
P. williamsoniana J.H.Willis R.P.J. de Kok 754 CANB AY883260
P. wudjariensis Orthia C.E. Woolcock 2250 CBG AY883261
Stonesiella Crisp & P.H.Weston
S. selaginoides (Hook.f.) Crisp & P.H.Weston R. Burns 258 CBG AF113791
Urodon Turcz.
U. capitatus Turcz. M.D. Crisp 8523 CANB AF113792
OUTGROUPS
Daviesia Sm.
D. elliptica Crisp M.D. Crisp 9051 CANB AF518130
D. flava Pedley I.R.D. Telford 12,054 CBG KY426177
D. longifolia Benth. M.D. Crisp 9246 CANB KY426196
Sphaerolobium Sm.
S. medium R.Br. M.D. Crisp 8942 CANB AF518136
S. minus Labill. M.D. Crisp 9054 CANB AF518135
S. nudiflorum (Meisn.) Benth. R. Butcher 1229A PERTH AF518137
3. Results
The sequence data comprised 200 sequences for trnL-F including the outgroup samples
(Table 2). Alignment and character statistics are shown in Table 3. Stationarity and mixing
were confirmed with an effective sample size of 79,932; a mean standard deviation of the
split allele frequencies of 4.931 ×10−3; and consistency in the tracer run.

Diversity 2021,13, 391 15 of 38
Table 3. Descriptive statistics for trnL-F alignment.
trnL-F
Number of samples 200
Sequence length range (bp) 518–934
Aligned length (bp) 934
GC content (%) 32.2
Variable sites 680
Gaps + missing characters (%) 11.6
Sphaerolobium is strongly supported as a monophyletic clade sister to core Mirbelieae.
The posterior tree is dominated by a hard polytomy from which all major clades of core
Mirbelieae are derived (Figure 1). This limits the degree to which relationships between
clades can be discussed, so instead, we will focus on the major clades that are strongly
supported within polytomy and their correlation with currently recognised genera. Sig-
nificantly, we recover a moderately supported clade that contains most Pultenaea species,
with four strongly supported subclades within this clade. Support for this clade, probably
the first to diverge in core Mirbelieae, appears to be influenced by outgroup selection, as
analyses including more outgroup taxa recover the same clade with strong support (>0.95)
(not shown). Three of these subclades are endemic to southeastern Australia, while one also
has numerous species in southwestern Australia. Core Oxylobium is strongly supported
as monophyletic, with the inclusion of Mirbelia oxylobiodes F.Muell., which is formally
transferred to Oxylobium below. Dillwynia,Eutaxia,Jacksonia,Leptosema and Phyllota are all
strongly supported as monophyletic based on only limited sampling. Aotus is separated
into two lineages based on just four included species, reflecting its diverse morphology.
Euchilopsis and Urodon are together weakly supported as sister to Phyllota. A clade of
Western Australian species currently included in Pultenaea s.l. is strongly supported as
sister to Latrobea, which is strongly supported as monophyletic based on two included
species. The position of ‘Otion microphyllum’ is not supported but it has morphological
affinities to Aotus.
Mirbelia is strongly supported as monophyletic, with the exception of M. oxylobioides
F.Muell. which clearly belongs in Oxylobium. The position of Pultenaea brachytropis Benth.
is not supported. It appears, on morphology, to be allied to P. craigiana C.F.Wilkins, Orthia
and Crisp (not sampled), and the two species may form an independent lineage. A second
clade of Western Australian species currently included in Pultenaea s.l. is strongly supported
as sister to a clade that includes Stonesiella,Almaleea and Pultenaea adunca Turcz., making
this a morphologically heterogenous clade. The relationship between the Chorizema species
and the type clade of Podolobium remains unclear, as these nodes are not supported in
our analyses, but it is possible that these two genera should be united. Three species of
Podolobium are strongly supported as sister to Callistachys lanceolata, and they are formally
recognised as Callistachys species below. Finally, there is strong support for the broad
concept of Gastrolobium adopted by Chandler et al. [
50
]. This definition of Gastrolobium
includes significant morphological diversity, reflecting the fact that it now includes the
former genera Brachysema R.Br., Jansonia Kippist and Nemcia Domin and Oxylobium lineare
(Benth.) Benth. (= Gastrolobium ebracteolatum G.Chandler and Crisp).
4. Discussion
As highlighted by Orthia et al. [
34
,
36
], the resolution of monophyletic genera is
impossible from these data unless the entirety of core Mirbelieae is incorporated in a
single mega-genus, to which the name Pultenaea would apply. Such a proposal has met
with significant resistance from both the general public and botanical community, so it
is our hope that recent advances in sequencing technology can generate novel data to
resolve relationships among the majority or all of the lineages that currently form the
hard polytomy recovered in our analyses. If these relationships can be resolved, then a
classification can be proposed that minimises taxonomic disruption and maintains as many

Diversity 2021,13, 391 16 of 38
traditionally recognised genera as possible, though significant re-circumscription of some
genera appears inevitable [34,36].
Careful assessment and reanalysis of molecular data (from the trnL-F marker) available
on GenBank, including a larger number of taxa, does recover a higher proportion of
supported monophyletic clades than found by Orthia et al. [
34
,
36
], including a clade
containing most Pultenaea species (130 of 148). Pultenaea is the largest genus in the tribe,
and it contains a high degree of morphological and phenotypic variation. The Pultenaea
species excluded from the core clade are all endemic to south-western Australia, and their
inclusion in Pultenaea has been questioned previously [96].
4.1. How to Resolve Generic Relationships within Mirbelieae?
The close evolutionary history of the Mirbelieae, which is potentially due to hybridisa-
tion, incomplete lineage sorting and recent radiation [
34
,
36
], can make genetic identification
of closely related species and individuals very difficult. This is often due to little genetic
sequence divergence, which is especially problematic when using traditional approaches
such as DNA barcoding [
151
]. However, for groups that may have recently diversified,
population genetic markers, such as microsatellites, are an excellent way to determine
genetic differences between closely related species [
152
–
154
]. In many cases, these closely
related species share a close geographic range, which can lead to interspecific hybridisation
and outbreeding depression, which can affect species delimitation [155].
Next-Generation Sequencing has revolutionised the way that we handle and think
about molecular data, due to its ability to sequence whole genomes. However, the costs and
bioinformatics processing associated with full-genome studies are often intractable under
most research budgets, and for answering most evolutionary and phylogenetic questions,
this quantity of data is not required [
156
]. Techniques, such as restriction associated
DNA sequencing (RADseq), offer an excellent and cost-effective approach that can be
used to find informative genome-wide markers and provide unprecedented phylogenetic
resolution [
157
]. However, markers developed by using RADseq are taxon-specific and not
cross-applicable between genera or families [
158
]. In recent years, huge advances have been
made in targeted sequence capture that allow for the enrichment of hundreds of informative
markers throughout the genome that are cross-applicable throughout angiosperms and
gymnosperms [159–163].
Johnson et al. [
164
] have recently developed a commercially available targeted baits
probe set (myBaits Angiosperms 353) that can generate a range of informative low-copy
exons from the nuclear genome, which are cross-applicable across angiosperms. This
baits set has particular utility for herbarium specimens that are up to 100 years old or
sometimes older [
162
]. To date, the Angiosperms 353 probe set has provided impressive
phylogenetic resolution in Schefflera J.R. Forst. and G. Forst. [
165
], Cyperaceae [
160
,
161
,
166
]
and Gesneriaceae [
167
], providing unprecedented support at both the backbone and species
levels. Additionally, the myBaits Angiosperms 353 kit, currently being used as part of the
Plant and Fungal Trees of Life (PAFTOL) [
163
,
164
,
168
], is a method that will form part of the
Phylogenomics Working Group (PWG) of the Genomics for Australian Plants Framework
Data Initiative (GAP; https://www.genomicsforaustralianplants.com (accessed on 25 April
2021)), setting a new standard for angiosperm genomics.
The resolution of tribal, generic and species relationships within Fabaceae tribe Mir-
belieae can realistically only be obtained by the targeted sampling of key species for the
generation of significant new molecular data [
34
,
36
]. The aim of such sampling is to develop
a new classification that minimises nomenclatural changes. Considering the complexity
and possible recent radiation of Mirbelieae, construction of a well-supported phylogeny
will require the use of rich and phylogenetically informative low-copy markers to allow
for competing phylogenetic signals in the data to be identified and resolved [
159
]. Using
targeted capture for Mirbelieae should not only provide good phylogenetic resolution of
genera but also at the species level [
169
]. Therefore, the use of target capture sequencing
should directly aid in species circumscription. A significant number of species complexes

Diversity 2021,13, 391 17 of 38
still require resolution (at least 28 in Pultenaea s. s.; unpubl. data by the authors), so this is
an important consideration when selecting a technique.
To accomplish this, we will utilise Next-Generation Sequencing in the form of targeted
baits capture using the myBaits Angiosperms 353 universal probe set in order to obtain
informative markers from the nuclear genome [
163
,
164
]. Enabling the sequencing of a
large number of informative low-copy markers throughout the nuclear genome at an
affordable cost should allow us to resolve the relationships between major clades and
therefore redefine genera. We will sample multiple individuals within the example species
complexes in order to test the utility of the PAFTOL markers [
163
] for resolution of such
taxonomic problems, guiding future studies. Where we are able to resolve species limits
within this project, we will revise or create new descriptions for the Australian eFlora and
publish new species as required.
A wide range of DNA samples have already been collected by the authors of this paper
and continued targeted fieldwork will ensure that the most important taxa and populations
can be included in order to address our key questions. Currently, silica-dried DNA samples
are held for ~95% of key taxa required for molecular work (subset from >1300 samples of
Faboideae held by project collaborators, with additional samples in the NSW DNA bank).
Further samples will be obtained through fieldwork for the remaining species to represent
all the key clades of Mirbelieae in order to provide the maximum phylogenetic resolution
and support from the Angiosperms 353 bait set [
162
,
163
]. Sampling from herbarium
specimens will also be undertaken where no field-collected material is available. It is
critical that all key taxa (including all type species) are included in a phylogeny to help
fully resolve generic level phylogenetic relationships in Mirbelieae.
4.2. Current Availability of Descriptive Information and Interactive Identification Tools
4.2.1. Online Floras
Most pea-flowered legumes are yet to be treated in the Flora of Australia, with the
first generic treatments only recently being added to the eFlora platform (e.g., Indigastrum
Jaub. & Spach.). To date, the only genus in Tribe Mirbelieae submitted for the Flora of
Australia is Gastrolobium (109 spp.). This treatment will be updated, where required, to
include new species (three putative new species and one published since submission [
84
])
for online publication.
Published revisions and regional flora treatments are available for most species in
15 Mirbelieae genera: Almaleea (5 spp.); Callistachys (5 spp.); Chorizema (28 spp.); Daviesia
(130 spp.); Dillwynia (32 spp.); Eutaxia (24 spp.); Gastrolobium (113 spp.); Gompholobium
(
45 spp.
); Jacksonia (74 spp.); Leptosema (12 spp.); Mirbelia (36 spp.); Pultenaea s.s. (130 spp.);
Sphaerolobium (22 spp.); Stonesiella (1 sp.); and Viminaria (1 sp.). Flora accounts of nine
genera without recent revisions still need to be completed: Aotus s.l. (22 spp.); Erichsenia
(1 sp.); Euchilopsis (1 sp.); Isotropis (15 spp.); Latrobea (9 spp.); ‘Otion’ (5 spp.); Oxylobium
(7 spp.); Phyllota (10 spp.); and Urodon (4 spp.). A few anomalous species may be placed
in the new genera, or in expanded definitions of existing genera. Two lineages currently
included in Pultenaea s.l., both endemic to Western Australia, appear to be well-supported
clades sister to other Western Australian genera, and new generic names may be required
for these clades if they continue to be supported by additional data and sampling. ‘Otion’
remains an informal name that also requires formal circumscription and validation. In
summary, current taxonomic descriptions are available in the published literature for
595 species
, while 160 species need revised or new descriptions (at least 32 of these species
are unnamed).
The release of profiles on the Australian eFlora under current generic circumscriptions
is not seen to be a viable option, as many of the generic and species boundaries are known
to be problematic. An example is the generic name ‘Otion’, first proposed in 1982, but never
formalised, leaving its constituent species orphaned. Combining these into Pultenaea s.l.
until generic boundaries are resolved would only further distort the circumscription of that
genus and be problematic. The definition of Pultenaea s.s. is also especially challenging

Diversity 2021,13, 391 18 of 38
at present, but the transfer of just 18 species from Pultenaea will likely resolve this as the
remainder of Pultenaea species are resolved as a monophyletic clade.
Published revisions are available for 595 species within Mirbelieae, leaving 160 species
that require revision, or new descriptions (Table 4). We aim for the completion of eFlora
treatments for all taxa in tribe Mirbelieae (24–33 genera and ca. 755 species). Treatments
have already been provided to ABRS for an additional 57 pea genera (including 483 species)
from other tribes. An ongoing project by R. Butcher, I.D. Cowie & T.D. Macfarlane et al.
([
170
–
176
]; Butcher, unpublished data) to revise Australian Tephrosia Pers. will add ca.
111 species
. At the completion of these projects, ca. 1370 species will be available for eFlora,
leaving a gap of ca. 393 species for completion of Fabaceae for the Flora of Australia.
Table 4.
Summary of genera in tribe Mirbelieae reflecting putative phylogenetic relationships, number with molecular data,
and progress towards eFlora treatments. Genera or clades marked with an * are yet to be adequately circumscribed and
require additional genetic data.
Genus
(or Unnamed Clade) Number of Species
No. of Species with
DNA Sequence
Data/No. of
Sequences
Published
Descriptions
Available
New or Revised
Descriptions Required
Core Mirbelieae
Almaleea 5 1/4 5 0
* Aotus s.s. 6 4/6 0 6
* Aotus Clade A 3 2/5 1 2
* Aotus Clade B 3 0 2 1
* Aotus Clade C 10 0 0 10
Callistachys s.l. 5 4/17 3 2
* Chorizema s.l. 28 18/32 28 0
* Dillwynia 36 3/7 4 32
Euchilopsis 1 1/4 0 1
* Eutaxia s.s. 12 3/6 12 0
Gastrolobium 109 82/313 109 0
Jacksonia 74 6/15 74 0
* Latrobea 9 4/12 3 6
Leptosema 12 2/5 12 0
* Mirbelia 26 15/65 28 8
* ’Otion’ ined. 5 1/2 1 4
Oxylobium 7 6/40 1 6
* Phyllota 10 2/7 1 9
* Podolobium 3 3/17 3 0
Pultenaea s.s. 130 58/222 83 47
* Pultenaea Clade A 1 1/3 1 0
* Pultenaea Clade B 2 2/6 2 0
* Pultenaea Clade C 9 9/28 7 2
* Pultenaea Clade D 6 6/18 6 0
* Sclerothamnus R.Br. 12 0 10 2
Stonesiella 1 1/1 1 0
Urodon 4 2/4 0 4
Mirbelieae s.l.
Daviesia 130 130/419 130 0
Erichsenia 1 1/3 0 1
Gompholobium 46 5/9 46 0
Isotropis 15 4/18 3 12
Sphaerolobium 22 3/7 22 0
Viminaria 1 1/4 1 0
Total Mirbelieae s.l. 755 385/1298 595 160

Diversity 2021,13, 391 19 of 38
4.2.2. Interactive Keys
The Pea Key provides a user-friendly resource for the identification of Australian
Faboideae [
138
]. However, technological changes since its first edition, released in 2007,
require the key to be updated with improved accessibility. Less than 50 species of Mirbelieae
need to be added and the functionality of the key must be improved in order to make it
more attractive and to increase its use by a wider audience. An expansion of the character
sets for Mirbelieae species would enable a more reliable tool in the identification of taxa to
species-level.
The Pea Key currently includes ca. 1500 species among 136 genera, based on
67 descriptive
characters (hosted by ANBG, last updated 2007). A new interactive key has recently been
published on FloraBase for Western Australian peas [
177
]. This key is based on 70 characters,
is well-illustrated, and is user-friendly. The WA key provides an excellent tool to aid in
the identification of 507 Mirbelieae species, and we aim to update existing data, images
and illustrations in The Pea Key for the 247 Mirbelieae species that occur outside of Western
Australia in order to align with, and complement, the Western Australian pea key. A
new interactive key has also been completed for Victoria, covering 309 species based on
50 characters [139].
Images and descriptive data were mostly absent in the first edition of The Pea Key.
Rather than creating a stand-alone dataset of descriptive information and images, it is
proposed that the second edition links to the newly created eFlora platform through unique
identifiers provided by the Atlas of Living Australia (ALA). Linking to the eFlora will
automatically utilise images in the Australian Plant Image Index (APII). Images of key
morphological features provide powerful identification aids and representative images
are provided to demonstrate the large range of morphological variation found in the tribe
(Figures 2–6).
4.3. Conclusions for Future Taxonomic Classification
Core Tribe Mirbelieae, as defined here, currently includes 18 genera, but our findings
show that at least some of these genera cannot be maintained in their current circumscrip-
tions. Options that create monophyletic genera include the recognition of a single large
genus, or up to 27 genera (in which case up to six genera would be new). We propose to use
targeted sampling of all recognised and possible novel genera, representing known diver-
sity in the group, in order to determine a novel generic level classification that minimises
taxonomic changes. We anticipate that with new data to confirm relationships between
the major lineages identified in our study, the majority of Pultenaea species can be retained
in that genus. The large genera Gastrolobium,Jacksonia and Mirbelia are likely to be main-
tained in the current circumscriptions. Most of the other genera in core Mirbelieae require
further sampling to determine revised monophyletic units that have utility as genera. This
resolution is critical to the appropriate placement of species in the Australian eFlora.

Diversity 2021,13, 391 20 of 38
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require further sampling to determine revised monophyletic units that have utility as gen-
era. This resolution is critical to the appropriate placement of species in the Australian
eFlora.
Figure 2. Morphological diversity of representative species from Fabaceae tribe Bossiaeeae (sister to Mirbelieae) and non-
core Mirbelieae. (A) Bossiaea eriocarpa. (B) Bossiaea preissii. (C) Bossiaea sericea. (D) Daviesia angulata. (E) Daviesia hakeoides.
(F) Daviesia mimosoides subsp. mimosoides. (G) Gompholobium knightianum. (H) Gompholobium marginatum. (I) Isotropis atro-
purpurea. (J) Isotropis cuneifolia subsp. cuneifolia (calyx and rear face of corolla). (K) Sphaerolobium drummondii. (L) Viminaria
Figure 2.
Morphological diversity of representative species from Fabaceae tribe Bossiaeeae (sister to Mirbelieae) and
non-core Mirbelieae. (
A
)Bossiaea eriocarpa. (
B
)Bossiaea preissii. (
C
)Bossiaea sericea. (
D
)Daviesia angulata. (
E
)Daviesia
hakeoides. (
F
)Daviesia mimosoides subsp. mimosoides. (
G
)Gompholobium knightianum. (
H
)Gompholobium marginatum.
(I)Isotropis atropurpurea.
(
J
)Isotropis cuneifolia subsp. cuneifolia (calyx and rear face of corolla). (
K
)Sphaerolobium drummondii.
(
L
)Viminaria juncea. Photographs by Russell Barrett except (
J
) by Kevin Thiele and (
L
) by Murray Fagg from the Australian
Plant Image Index: http://www.anbg.gov.au/photo (accessed on 11 June 2021)).

Diversity 2021,13, 391 21 of 38
Diversity 2021, 13, x FOR PEER REVIEW 21 of 39
juncea. Photographs by Russell Barrett except (J) by Kevin Thiele and (L) by Murray Fagg from the Australian Plant Image
Index: http://www.anbg.gov.au/photo (accessed on 11 June 2021)).
Figure 3. Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae). (A) Almaleea
incurva. (B) Aotus gracillima. (C) Aotus lanigera. (D) Callistachys lanceolata. (E) Callistachys scandens. (F) Chorizema rhombeum.
(G) Chorizema varium. (H) Dillwynia ericifolia. (I) Dillwynia phylicoides. (J) Dillwynia retorta. (K) Dillwynia sericea. L. Erichsenia
uncinata. Photographs by Russell Barrett except (A,D,G) by Murray Fagg and (B,L) by Kevin Thiele ((A,B,D,G,L) from the
Australian Plant Image Index: http://www.anbg.gov.au/photo (accessed on 11 June 2021)).
Figure 3. Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae). (A)Almaleea
incurva. (
B
)Aotus gracillima. (
C
)Aotus lanigera. (
D
)Callistachys lanceolata. (
E
)Callistachys scandens. (
F
)Chorizema rhombeum.
(
G
)Chorizema varium. (
H
)Dillwynia ericifolia. (
I
)Dillwynia phylicoides. (
J
)Dillwynia retorta. (
K
)Dillwynia sericea. (
L
)Erichsenia
uncinata. Photographs by Russell Barrett except (
A
,
D
,
G
) by Murray Fagg and (
B
,
L
) by Kevin Thiele ((
A
,
B
,
D
,
G
,
L
) from the
Australian Plant Image Index: http://www.anbg.gov.au/photo (accessed on 11 June 2021)).

Diversity 2021,13, 391 22 of 38
Diversity 2021, 13, x FOR PEER REVIEW 22 of 39
Figure 4. Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae). (A,B)
Euchilopsis linearis. (C) Eutaxia microphylla. (D) Eutaxia virgata. (E) Gastrolobium hookeri. (F) Gastrolobium propinquum. (G)
Gastrolobium reticulatum. (H) Gastrolobium sericeum. (I) Gastrolobium sowardii. (J) Jacksonia aculeata. (K) Jacksonia capitata. (L)
Jacksonia compressa. Photographs by Russell Barrett except (C), by Murray Fagg ((C) from the Australian Plant Image Index:
http://www.anbg.gov.au/photo (accessed on 11 June 2021)).
Figure 4.
Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae).
(A,B)Euchilopsis linearis.
(
C
)Eutaxia microphylla. (
D
)Eutaxia virgata. (
E
)Gastrolobium hookeri. (
F
)Gastrolobium propin-
quum. (
G
)Gastrolobium reticulatum. (
H
)Gastrolobium sericeum. (
I
)Gastrolobium sowardii. (
J
)Jacksonia aculeata. (
K
)Jacksonia
capitata. (
L
)Jacksonia compressa. Photographs by Russell Barrett except (
C
), by Murray Fagg ((
C
) from the Australian Plant
Image Index: http://www.anbg.gov.au/photo (accessed on 11 June 2021)).

Diversity 2021,13, 391 23 of 38
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Figure 5. Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae). (A) Jacksonia
restioides. (B) Latrobea glabrescens. (C) Leptosema anomalum. (D) Leptosema aphyllum. (E) Leptosema chambersii. (F) Mirbelia
dilatata. (G) Mirbelia floribunda. (H) Mirbelia rhagodioides. (I) Mirbelia rubiifolia. (J) Mirbelia trichocalyx. (K) Mirbelia viminalis.
(L) ‘Otion’ simplicifolium. Photographs by Russell Barrett except (A) by Michael Crisp, (D,E) by Murray Fagg and (F) by
Kevin Thiele (D–F) from the Australian Plant Image Index: http://www.anbg.gov.au/photo (accessed on 11 June 2021)).
Figure 5.
Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae). (
A
)Jacksonia
restioides. (
B
)Latrobea glabrescens. (
C
)Leptosema anomalum. (
D
)Leptosema aphyllum. (
E
)Leptosema chambersii. (
F
)Mirbelia
dilatata. (
G
)Mirbelia floribunda. (
H
)Mirbelia rhagodioides. (
I
)Mirbelia rubiifolia. (
J
)Mirbelia trichocalyx. (
K
)Mirbelia viminalis.
(
L
)‘Otion’ simplicifolium. Photographs by Russell Barrett except (
A
) by Michael Crisp, (
D
,
E
) by Murray Fagg and (
F
) by
Kevin Thiele (D–F) from the Australian Plant Image Index: http://www.anbg.gov.au/photo (accessed on 11 June 2021)).

Diversity 2021,13, 391 24 of 38
Diversity 2021, 13, x FOR PEER REVIEW 24 of 39
Figure 6. Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae). (A) Oxy-
lobium cordifolium. (B) Oxylobium oxylobioides (flowering). (C) Oxylobium oxylobioides (fruiting). (D) Podolobium acicuiliferum.
(E) Pultenaea brachyphylla. (F) Pultenaea ferruginea. (G) Pultenaea maritima. (H) Pultenaea procumbens. (I) Pultenaea sp. Olinda.
(J) Pultenaea stipularis. (K) Stonesiella selaginoides. (L) Urodon dasyphyllus. Photographs by Russell Barrett except (A,D,K,L)
by Murray Fagg ((A,D,K,L) from the Australian Plant Image Index: http://www.anbg.gov.au/photo (accessed on 11 June
2021)).
Figure 6.
Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae). (
A
)Oxy-
lobium cordifolium. (
B
)Oxylobium oxylobioides (flowering). (
C
)Oxylobium oxylobioides (fruiting). (
D
)Podolobium acicuil-
iferum.
(E)Pultenaea brachyphylla.
(
F
)Pultenaea ferruginea. (
G
)Pultenaea maritima. (
H
)Pultenaea procumbens. (
I
)Pultenaea sp.
Olinda. (
J
)Pultenaea stipularis. (
K
)Stonesiella selaginoides. (
L
)Urodon dasyphyllus. Photographs by Russell Barrett except
(
A
,
D
,
K
,
L
) by Murray Fagg ((
A
,
D
,
K
,
L
) from the Australian Plant Image Index: http://www.anbg.gov.au/photo (accessed
on 11 June 2021)).

Diversity 2021,13, 391 25 of 38
Author Contributions:
Conceptualization, R.L.B. and J.A.R.C.; methodology, R.L.B.; software, R.L.B.,
J.A.R.C., M.A.M.R.; writing—original draft preparation, R.L.B., J.A.R.C., L.G.C., M.D.C., P.C.J., B.J.L.,
M.A.M.R. and P.H.W.; writing—review and editing, R.L.B., J.A.R.C., L.G.C., M.D.C., P.C.J., B.J.L.,
M.A.M.R. and P.H.W. All authors have read and agreed to the published version of the manuscript.
Funding:
This research was funded by a Postdoctoral Fellowship Grant from the Australian Biological
Resources Study (ABRS) National Taxonomy Research Grant Program (NTRGP 4-EHP5TK3) to James
Clugston and collaborators who are all co-authors of this paper.
Data Availability Statement:
All data used in this study are available at https://www.ncbi.nlm.nih.
gov/ (accessed on 25 April 2021).
Acknowledgments:
We thank the many people who have assisted us with fieldwork, collecting
specimens. The logistical support of Greg Harper on long field trips to the remotest parts of Australia
with Lyn Cook is particularly acknowledged. Colleagues have provoked many thoughtful discussions
that have all served to improve our knowledge of this wonderful group of peas. We particularly thank
Jim Ross as coordinator of the 150 Conference in Melbourne in 2003 which first bought the present
authors together in discussion on this topic through a session on generic concepts in the Australian
flora. Photographs by Murray Fagg and Kevin Thiele are reproduced from the Australian Plant
Image Index: http://www.anbg.gov.au/photo (accessed on 11 June 2021). Constructive comments
from two reviewers and handling editor Ashley Egan improved the manuscript.
Conflicts of Interest: The authors declare no conflict of interest.
Appendix A
Taxonomy
Mirbelieae s.s.
While many changes in generic circumscription are expected, most require more data
and better sampling before the best nomenclatural solutions can be determined [
34
,
36
].
However, there is sufficient support to justify a few taxonomic changes and these are
validated below. The opportunity is taken to select types for genera where type species
have never been designated, as far as we can determine. Specific lectotypes for species and
infraspecific taxa are also designated in order to place names correctly in advance of a new
global checklist of Fabaceae (R. Govaerts et al. in prep. http://sftp.kew.org/pub/data_
collaborations/Fabaceae/DwCA/ (accessed on 29 June 2021)). A nomenclatural summary
is presented for all Mirbelieae genera.
Almaleea
Crisp & P.H. Weston, Telopea 4(2): 309 (1991). Type:Almaleea incurvata (A. Cunn.)
Crisp & P.H. Weston
Aotus
Sm. in K.D.E. Koenig, & J. Sims (eds), Annals of Botany 1(3): 504 (1805). Type:Aotus
villosa (Andrews) Sm. [=A. ericoides (Vent.) D.Don]
Callistachys
Vent., Jardin de la Malmaison 2(20): 115, pl. 115 (1805). Type:Callistachys
lanceolata Vent.
Notes:Callistachys has been rejected against the conserved name Oxylobium Andrews
(1807) [
178
] if the two genera are united, but this has no effect when they are considered
distinct genera. We anticipate that the two genera will both remain accepted.
Three eastern Australian species previously included in Podolobium are clearly closely
related to the Western Australian species Callistachys lanceolata (Figure 1C; [
179
]), and they
are here included in Callistachys where names are already available [40].
Callistachys alpestris (F. Muell.) Kuntze, Revisio Generum Plantarum 1: 168 (1891).
Oxylobium alpestre F. Muell., Definitions of rare or hitherto undescribed Australian plants 5
(1855). Podolobium alpestre (F. Muell.) Crisp & P.H. Weston in Crisp, M.D. & Doyle, J.J. (eds.)
Advances in Legume Systematics 7: 280 (1995).
Type citation: ‘Not unfrequent in the higher parts of the Australian Alps.’
Type: Mount Buller, 5000 ft [1540 m], Victoria, Apr. 1853, F. Mueller s.n. (lecto, here
designated: MEL 624821; isolecto: K 000642469). Residual syntypes: Australian Alps,
Victoria, F. Mueller s.n. (K 000642470, MEL 624820). Mount Timbertop, Victoria, 26 Mar.

Diversity 2021,13, 391 26 of 38
1853, F. Mueller s.n. (MEL 624824). Subalpine mountains on the Berrima, Victoria, F. Mueller
s.n. (MEL 624822). In alpestre Great Dividing Range and in alpe Mt Cobberas, Victoria,
5-6000 ft, Jan. 1854, F. Mueller s.n. (K 000642467, MEL 624823, TCD 0014295*). Mitta
Mitta, 4-5000 ft, F. Mueller s.n. (K 000642468). Munyang Mountains, Victoria, F. Mueller s.n.
(BM 000885480, M 0219088*, PH 00017792*). Mount Hotham, Victoria, F. Mueller s.n. (BM
000885479). Victorian Alps, Walter s.n. (BM 000885478).
Oxylobium alpestre F. Muell., Victoria—Parliamentary Papers- Votes and Proceedings of the
Legislative Assembly 3, 12 (1853), nom. inval., nom. nud.
Notes: Mueller [
180
] described the fruit of this species, with no details of the flowers. Nu-
merous syntypes have been located, many without collection dates. It is unknown whether
he did have flowering material, so both fruiting and flowering sheets are considered syn-
types. We select a fruiting sheet at MEL as the lectotype as this is one of the the largest
specimens at MEL, it has a collection date firmly establishing that it was collected before
the species was named, and it is also a good match for the protologue. There is also a
duplicate at K.
Callistachys procumbens (F. Muell.) Kuntze, Revisio Generum Plantarum 1: 168 (1891).
Oxylobium procumbens F. Muell., Definitions of rare or hitherto undescribed Australian plants
4–5 (1855). Podolobium procumbens (F. Muell.) F. Muell. ex Crisp & P.H. Weston in Crisp,
M.D. & Doyle, J.J. (eds.) Advances in Legume Systematics 7: 281 (1995).
Type citation: ‘On wooded hills; for instance, at Mount Disappointment, in the Goulburn
Ranges, on the Delatite, in the Black Forest, at Ballarat, etc.’
Type: Black Forest, Victoria, Dec. 1852, F. Mueller s.n. (lecto, here designated: MEL 624827;
isolecto: MEL 624828). Residual syntypes: Delatite River, Victoria, 18 Mar. 1853, F. Mueller
s.n. (K 000642444, MEL 624829). Gipps Land [Gippsland] Range, F. Mueller s.n. (TCD
0014277). In the stringybark ranges ... the Glenelg and Goulbourn [Rivers] Victoria, 8 Feb.
1853, F. Mueller s.n. (MEL 569724). Australia Felix [Victoria], n.d., F. Mueller s.n. (MEL
624825). Possible syntype: Hume River, New South Wales, [received 1887], F. Mueller s.n. (M
0219079*).
Podolobium procumbens F. Muell., Victoria—Parliamentary Papers- Votes and Proceedings of the
Legislative Assembly 3, 12 (1853), nom. inval., nom. nud.
Notes: Mueller [
180
] cited a range of localities. We select a specimen at MEL from the
Delatite River as lectotype, as it is excellent fertile material and there is a duplicate sheet. A
specimen from the Hume River may or may not fall within the scope of the protologue and
has no collection date, so its type status is uncertain.
Callistachys scandens (Sm.) Kuntze, Revisio Generum Plantarum 1: 168 (1891).
Chorizema scandens Sm. in K.D.E. Koenig, & J. Sims (eds), Annals of Botany 1(3): 506 (1805).
Podolobium scandens (Sm.) DC., Prodromus Systematis Naturalis Regni Vegetabilis 2: 103 (1825).
Oxylobium scandens (Sm.) Benth., Commentationes de Leguminosarum Generibus 6 (1837).
Type citation: ‘Port Jackson.’
Type: Port Jackson [Sydney], New South Wales, 1793, Mr. White s.n. (lecto, designated by
M.D. Crisp et al. in Crisp, M.D. & Doyle, J.J. (eds), Advances in Legume Sytematics 7: 281
(1995)): LINN-HS728-3; isolecto: ?BM 000885551, LIV, P 00337501).
Note: See Crisp & Weston [
40
] for a full list of synonyms. Crisp et al. [
33
] cite the type of
the name Chorizema scandens as ‘Holo: Port Jackson, N.S. Wales, Mr. White, 1793 (LINN);
iso: ?BM, LIV, ?P.’ This is here treated as effective lectotypification in accordance with ICN
Art. 7.11 [181].
Chorizema
Labill., Relation du Voyage a la Recherche de la Perouse 1: 404, pl. 21 (1800). Type:
Chorizema ilicifolia Labill.
Orthotropis Benth. ex Lindl., Edwards’s Botanical Register Appendix: xvi (1839). Type:Or-
thotropis pungens Lindl. [=Chorizema aciculare (DC.) C.A. Gardner]
Note:Podolobium s.s. may be embedded within Chorizema, but further data are required to
test this.

Diversity 2021,13, 391 27 of 38
Dillwynia
Sm. in K.D.E. Koenig, & J. Sims (eds.), Annals of Botany 1(3): 510 (1805), non
Roth (1806). Type:Dillwynia ericifolia Sm., designated by L.K.G. Pfeiffer, Nomencl. Bot. 12:
136 (1874).
Dillwynia sect. Xeropetalum R.Br. ex Sims, Curtis’s Botanical Magazine 48: 2247, subt. (1821).
Type:Dillwynia cinerascens R.Br.
Xeropetalum Rchb., Conspectus regni vegetabilis per gradus naturales evoluti. Tentamen 154
(1828), nom. inval., nom. nud., non Delile (1826).
Dillwynia sericea subsp. glabriflora (Blakely) Jobson & P.H. Weston, comb. et stat. nov.
Basionym:Dillwynia sericea var. glabriflora Blakely, The Australian Naturalist 10(6): 185 (1939).
Type: Pilliga Forest [Scrub], Sept. 1913, E.H.F. Swain 29 (holo: NSW 40780; iso: CBG
8313107).
Dillwynia sericea subsp. rudis (Sieber ex DC.) Jobson & P.H. Weston, comb. et stat. nov.
Basionym:Dillwynia rudis Sieber ex DC., Prodromus Systematis Naturalis Regni Vegetabilis
2: 109 (1825). Type: [Sydney region, New South Wales], 1823, F. Sieber pl. exs. nov.-holl. n.
400 (lecto, here designated: G 00488223*; isolecto: BM 000885863, BR 0000013455989*,
G 00365064*, G 00365100*, H 1275852*, K 000858585, M 0219096*, MEL 624489, MO
277045*, MPU 021245*, NSW 40727, S-G-9090, TCD 0014717*, W 0045220, W 0045221,
W 19890005724).
Notes: There are many sheets of Sieber’s pl. exs. nov.-holl. n. 400, including at G, so we here
designate the sheet from de Candolle’s herbarium as lectotype as we know this is material
originally examined by de Candolle.
Dillwynia sparsifolia (F. Muell.) Jobson & P.H. Weston, comb. nov.
Basionym:Eutaxia sparsifolia F. Muell., Definitions of rare or hitherto undescribed Australian
plants 39–40 (1855).
Type citation: ‘In the desert scrub towards the mouth of the Murray River. Found also at
Tumbay Bay by Mr. C. Wilhelmi.’
Type: Tumby Bay, Spencers Gulf, [South Australia], C. Wilhelmi s.n. (lecto, here designated:
MEL 2138746). Probable syntype: Near Lake Alexandrina, [South Australia], Oct. 1848, F.
Mueller s.n. (MEL 2138744).
Notes: Mueller ’s concept of Eutaxia sparsifolia appears to equate with Eutaxia patula F. Muell.
ex D. Dietr [
180
]. The material cited for Eutaxia sparsifolia from ‘towards the mouth of the
Murray River’ is probably the MEL sheet also identified as a type of Eutaxia patula, but it
does not have the name E. sparsifolia on the sheet. We therefore designate the Wilhelmi
specimen at MEL as the lectotype of as the only definite original material of E. sparsifolia
located.
Dillwynia trichopoda (Blakely) Jobson & P.H. Weston, comb. et stat. nov.
Basionym:Dillwynia parvifolia var. trichopoda Blakely, The Australian Naturalist 10(5): 162
(1939).
Type: Hill Top, Southern Tableland, [New South Wales], Sept. 1899, J.H. Maiden & J.L.
Boorman s.n. (lecto, here designated: NSW 40290; isolecto: CBG 8313090, NSW450238).
Notes: Three sheets of the single gathering cited in the protologue have been located,
including one sheet at CBG (ex NSW). We here designate the sheet with the largest amount
of material as the lectotype.
Euchilopsis
F. Muell., The Chemist and Druggist with Australasian Supplement 5(13): 13 (1882).
Type:Euchilopsis linearis (Benth.) F. Muell.
Sphaerolobium sect. Euchiloides Benth., Flora Australiensis 2: 63, 67 (1864). Euchilodes Kuntze
in T. von Post & O. Kuntze, Lexikon Generum Phanerogamarum 212 (1903), nom. illeg., nom.
superfl.Type:Sphaerolobium euchilus Benth., nom. illeg., nom. superfl. [=Euchilopsis linearis
(Benth.) F. Muell.]
Eutaxia
R.Br. in W.T. Aiton, Hortus Kewensis Edn. 2, 3: 16 (1811). Type:Eutaxia myrtifolia
(Sm.) R.Br., nom. illeg., designated by C.F. Wilkins et al., Nuytsia 20: 111 (2010).
Sclerothamnus R.Br. in W.T. Aiton, Hortus Kewensis Edn. 2, 3: 16 (1811). Eutaxia sect.
Sclerothamnus (R.Br.) F. Muell. Fragmenta Phytographiae Australiae 1(1): 7 (1858). Type:
Sclerothamnus microphyllus R.Br. [= Eutaxia microphylla (R.Br.) C.H. Wright & Dewar]

Diversity 2021,13, 391 28 of 38
Gastrolobium
R.Br. in W.T. Aiton, Hortus Kewensis Edn. 2, 3: 16 (1811). Type:Gastrolobium
bilobum R.Br.
Brachysema R.Br. in W.T. Aiton, Hortus Kewensis Edn. 2, 3: 10 (1811). Type:Brachysema
latifolium R.Br. [= Gastrolobium latifolium (R.Br.) G.Chandler & Crisp]
Brachysema sect. Eubrachysema Benth., Flora Australiensis 2: 9, 10 (1864), nom. inval.
Jansonia Kippist, The Gardeners’ Chronicle and Agricultural Gazette 19: 307 (1847). Type:
Jansonia formosa Kippist ex Lindl. [= Gastrolobium formosum (Kippist ex Lindl.) G.Chandler
& Crisp]
Cryptosema Meisn. in J.G.C. Lehmann (ed.), Plantae Preissianae 2(2-3): 206 (1848). Type:
Cryptosema pimeleoides Meisn. [= Gastrolobium formosum (Kippist ex Lindl.) G.Chandler &
Crisp]
Nemcia Domin, Preslia 2: 27 (1923). Type:Nemcia coriacea (Sm.) Domin [= Gastrolobium
coriaceum (Sm.) G. Chandler & Crisp]
Notes: Domin included twelve species in his new genus Nemcia [
182
]. We here select N.
coriacea as the type species, as its phylogenetic position is strongly supported, and it is a
well-defined species that is representative of the genus sensu Domin [182].
Cupulanthus Hutch., Dicotyledones.The Genera of Flowering Plants (Angiospermae) 1: 341
(1964). Type: not designated.
Jacksonia
R.Br. ex Sm. in A. Rees (ed.), The Cyclopaedia 18: - (1811). Type:Jacksonia spinosa
(Labill.) R.Br. ex Sm., designated by J.A. Chappill et al., Australian Systematic Botany 20(6):
476 (2007).
Piptomeris Turcz., Bulletin de la Societe Imperiale des Naturalistes de Moscou 26(1): 257 (1853).
Jacksonia sect. Piptomeris (Turcz.) Kuntze in T. von Post & O. Kuntze, Lexikon Generum
Phanerogamarum 294 (1903). Type:Piptomeris aphylla Turcz. [= Jacksonia racemosa Meisn.]
Latrobea
Meisn. in J.G.C. Lehmann (ed.), Plantae Preissianae 2(2-3): 219 (1848). Type:Latrobea
genistioides (Meisn.) Meisn.
Latrobea sect. Eulatrobea Benth., Flora Australiensis 2: 140 (1864), nom. inval.
Leptocytisus Meisn. in J.G.C. Lehmann (ed.), Plantae Preissianae 2(2-3): 211 (1848). Latrobea
sect. Leptocytisis (Meisn.) Benth., Flora Australiensis 2: 140, 141 (1864). Type:Leptocytisus
tenellus (Meisn.) Meisn. [= Latrobea tenella (Meisn.) Benth.]
Notes: Meissner included two species in his new genus Latrobea, both from south-west
Western Australia [
65
]. We here select L. genistioides as the type species as it is well
represented by type material, while L. brunonis (Benth.) Meisn. is not.
Leptosema
Benth., Commentationes de Leguminosarum Generibus 20 (1837). Type:Leptosema
bossiaeoides Benth.
Kaleniczenkia Turcz., Bulletin de la Societe Imperiale des Naturalistes de Moscou 26(1): 252 (1853).
Type:Kaleniczenkia daviesioides Turcz. [= Leptosema daviesioides (Turcz.) Crisp]
Burgesia F. Muell., Fragmenta Phytographiae Australiae 1(10): 222 (1859). Type:Burgesia
homaloclada F. Muell. [= Leptosema aphyllum (Hook.) Crisp]
Mirbelia
Sm. in K.D.E. Koenig, & J. Sims (eds), Annals of Botany 1(3): 511 (1805). Type:
Mirbelia reticulata Sm.
Dichosema Benth. in S.F.L. Endlicher et al., Enumeratio plantarum quas in Novae Hollandiae
ora austro-occidentali ad fluvium Cygnorum et in Sinu Regis Georgii collegit Carolus liber baro
de Hügel 35 (1837). Mirbelia sect. Dichosema (Benth.) Kuntze in T. von Post & O. Kuntze,
Lexikon Generum Phanerogamarum 368 (1903). Type:Dichosema spinosum Benth. [= Mirbelia
spinosa (Benth.) Benth.]
Oxycladium F. Muell. in W.J. Hooker (ed.), Hooker’s Journal of Botany and Kew Garden
Miscellany 9: 20 (1857). Type:Oxycladium semiseptatum F. Muell. [= Mirbelia viminalis (A.
Cunn. ex Benth.) C.A. Gardner]
Oxylobium
Andrews, The Botanist’s Repository for New, and Rare Plants 7(101): t. 492 (1807),
nom. cons.Type:Oxylobium cordifolium Andrews
Oxylobium oxylobioides (F. Muell.) Crisp & R.L. Barrett, comb. nov.
Basionym:Mirbelia oxylobioides F. Muell., Fragmenta Phytographiae Australiae 2(16): 154 (1861).

Diversity 2021,13, 391 29 of 38
Type citation: ‘In vallibus profundis rupestribus montium Haidinger Range as originem
fluviorum Macallister et Mitchell River, altitudine 3000–4000.’
Type: Sources of the Macallister [Macalister] River, Mar. 1861, F. Mueller s.n. (lecto, here
designated: MEL 624816). Possible syntypes: Mc’Allister [Macalister] River, no date, F.
Mueller s.n. (MEL 624817; NY 00026421). Mount Legar and Range to the N.W., no date, F.
Mueller s.n. (K 000642571).
Typification: While MEL 624815 is labelled as a type specimen, it is clearly labelled as
collected in 1863, two years after the species was named, so it cannot be original material.
There is a duplicate of this specimen at NSW (NSW 31714) which is also dated 1863. The
later collection locality near the Macalister River creates doubt over the undated collections
that are simply labelled ‘Mc’Allister River’ as to whether or not they are syntypes [Muller’s
duplicate labels were often simplified from the original label]. Only the specimen here
designated as lectotype can be confidently considered original material.
Notes: This species consistently groups with Oxylobium in phylogenetic analyses and
notably it only has an abaxial septum in the longitudinally divided pod, a character shared
with O. robustum Joy Thomps. [39,47].
Phyllota
(DC.) DC. ex Benth. in S.F.L. Endlicher et al., Enumeratio plantarum quas in Novae
Hollandiae ora austro-occidentali ad fluvium Cygnorum et in Sinu Regis Georgii collegit Carolus
liber baro de Hügel 33 (1837). Pultenaea sect. Phyllota DC. in A.P. de Candolle (ed.), Prodromus
Systematis Naturalis Regni Vegetabilis 2: 113 (1825). Type:Pultenaea phylicoides Sieber ex DC.
[= Phyllota phylicoides (Sieber ex DC.) Benth.]
Walpersia Harv. in Harvey & Sonder, Fl. Cap. 2: 26 (1861), nom. cons., non Reissek ex Endl.
(1840). Type:Walpersia burtonioides Harv. [= Phyllota squarrosa (Sieber ex DC.) Benth.]
Notes: de Candolle included four species of Pultenaea in his section Phyllota:P. aspera Sieber
ex DC., P. comosa Sieber ex DC., P. squarrosa Sieber ex DC. and P. phylicoides Sieber ex DC., all
collected by Franz Sieber in the vicinity of Sydney, New South Wales [
57
]. All four species
were recognised by Bentham [
59
] when he raised the section to genus rank. However,
Bentham later reduced three of these to synonymy under Phyllota phylicoides (Sieber ex DC.)
Benth. [62]Phyllota squarrosa (Sieber ex DC.) Benth. has since been reinstated.
We note that Bentham [
59
] specifically attributed the genus name to de Candolle (probably
based on de Candolle ([
57
]: 113) stating ‘An genus proprium?’ [
59
] We therefore accept the
authorship of the genus name as ‘(DC.) DC. ex Benth.’
Jancey [
92
] undertook a detailed study of Phyllota in New South Wales, providing detailed
notes on the type species for the species, but did not select a type for the genus. We here
select Pultenaea phylicoides as the type of Pultenaea sect. Phyllota (and therefore of Phyllota),
as it is only one of the four original species later recognised by Bentham [62].
Walpersia burtonioides was described as a new genus and species from South Africa based
on erroneous label data and it is actually an Australian taxon [183].
Phyllota barbata
Benth. in S.F.L. Endlicher et al., Enumeratio plantarum quas in Novae
Hollandiae ora austro-occidentali ad fluvium Cygnorum et in Sinu Regis Georgii collegit Carolus
liber baro de Hügel 33 (1837).
Type citation: ‘King Georges Sound (Hügel.)’
Type: King Georges Sound, Western Australia, C.A.A. von Hügel 97 (lecto, here designated:
W 0046869). Residual syntypes: King Georges Sound, Western Australia, C.A.A. von Hügel 2
(W 0046868); King Georges Sound, Western Australia, C.A.A. von Hügel 3 (W 0046868).
Phyllota villosa Turcz., Bull. Soc. Imp. Naturalistes Moscou 26(I): 267 (1853).
Type: Western Australia, 1842, J. Gilbert 255 (lecto, here designated: KW 001001187; isolecto:
K 000858466).
Typification: There are three sheets of Phyllota barbata at W collected by von Hügel, each
with different collection numbers, so they are here regarded as syntypes. W 0046869 is
designated as the lectotype as it is the only sheet that bears the name ‘Phyllota barbata’ on
the original labels.

Diversity 2021,13, 391 30 of 38
Two sheets of the type collection of Phyllota villosa have been located. We here designate
the sheet at KW as lectotype as this is the sheet used by Turczaninow when describing the
species.
Notes: Orthia et al. [
96
] noted the morphological similarity of Phyllota barbata and Pultenaea
barbata and even suggested they were possibly conspecific. We conclude that the two names
might best be treated as a single species based on study of the type specimens, however the
names are currently both in use for two quite distinct taxa in Western Australia, so further
studies are advised on the application of both names. The two taxa were named in separate
genera, but independently given the same species epithet. If the two taxa are united, the
younger name has priority if the taxon is included in Pultenaea (as Orthia et al. [
96
] did);
however, in Phyllota the older name has priority.
Podolobium R.Br. in W.T. Aiton, Hortus Kewensis Edn 2, 3: 9 (1811).
Type:Podolobium trilobatum R.Br. [= P. ilicifolium (Andrews) Crisp & P.H. Weston; =Chorizema
trilobum Sm., non Chorizema ilicifolium Labill. (1800).]
Note:Podolobium is not monophyletic as circumscribed in recent literature (e.g., [
40
,
134
]),
however, it probably is monophyletic with the transfer above of three species to Callistachys.
The type species of Podolobium may be resolved within Chorizema which would have priority
if the two genera are combined.
Pultenaea
Sm., A Specimen of the Botany of New Holland 1(3): 35 (1794). Type:Pultenaea
stipularis Sm.
Euchilus R.Br. in W.T. Aiton, Hortus Kewensis Edn. 2, 3: 17 (1811). Pultenaea sect. Euchilus
(R.Br.) F. Muell., Fragmenta Phytographiae Australiae 1(1): 8 (1858). Type:Euchilus obcordatus
R.Br. [= Pultenaea heterochila F. Muell.]
Bartlingia Brongn., Memoire sur la famille des Rhamnees 66 (1826), nom. illeg., non Reichb.
(1824). Type:Bartlingia obovata (DC.) Meisn.
Spadostyles Benth., Commentationes de Leguminosarum Generibus 16 (1837). Type:Spadostyles
cunninghamii Benth. [=Pultenaea spinosa (DC.) H.B. Will.]
Notes: We here designate Spadostyles cunninghamii Benth. as the type species of Spadostyles
Benth., as this is the best described of the two species included by Bentham [
60
], and the
only name that is legitimate.
Pultenaea barbata
C.R.P. Andrews, Journal of the West Australian Natural History Society
2(1): 38–39 (1904).
Type citation: ‘I found this species in flower near the Phillips River in Oct. 1903.’
Type: Phillips River, Western Australia, Oct. 1903, C.R.P. Andrews s.n. (lecto, here designated:
PERTH 01025805; isolecto: NSW 36498).
Pultenaea andrewsii W.E. Blackall & B. Grieve, How to know Western Australian wildflowers: a
key to the flora of the temperate regions of Western Australia 1: 234, (1954), nom. inval., nom. nud.
Typification: There are two sheets of the type collection of Pultenaea barbata. We here
designate the PERTH sheet as the lectotype as it can be expected that both sheets were held
in Western Australia (prior to the formal establishment of a state herbarium) at the time of
description, with the subsequent distribution of one sheet to NSW.
Notes: See discussion under Phyllota barbata above.
Pultenaea benthamii
F. Muell., Definitions of rare or hitherto undescribed Australian plants 5
(1855).
Type: amongst rocks on the top of Mount Abrupt, F. Mueller s.n. (lecto, designated by R.P.J.
de Kok & J.G. West, Australian Systematic Botany 17(3): 276 (2004), K 000118882); isolecto:
BM 000544575, MEL 567120, TCD 0014523).
Pultenaea benthamii var. elatior Benth. Fl. Austral. 2: 114 (1864).
Type citation: ‘Yowaka river, and foot of mount William, F. Mueller.’
Type: Yowaka River, New South Wales, Apr. 1860, F. Mueller s.n. (lecto, here designated: K
000119047; isolecto: MEL 567122). Residual syntypes: Foot of Mount William, Nov. 1853,
F. Mueller s.n. (K; MEL 567121, MEL 567123, ?MEL 627862) [= Pultenaea humilis Benth. ex
Hook.f.].

Diversity 2021,13, 391 31 of 38
Notes: de Kok & West [
93
] noted that var. elatior was based on two collections representing
two taxa. We here choose a lectotype which places the variety as a synonym of P. benthamii,
while the residual syntype represents P. humilis.
Pultenaea elusa (J.D. Briggs & Crisp) R.L. Barrett & Clugston, comb. et stat. nov.
Basionym: Pultenaea parrisiae subsp. elusa J.D. Briggs & Crisp, Telopea 5: 652 (1994).
Type: New South Wales: Central Tablelands: Wingello in swamps, 30 Sep 1938, W.F. Blakely
s.n. (holo: NSW 38321).
Pultenaea elusa de Kok & J.G. West, Austral. Syst. Bot. 17: 288 (2004), nom. inval.
Note: de Kok and West [
93
] attempted to raise Pultenaea parrisiae subsp. elusa to the species
rank, however they unfortunately cited the entire page range of the publication, so the
combination was not validly published (ICN Article 41.5 [
181
]). We therefore provide a
valid combination here.
Pultenaea recurvifolia
(Benth.) H.B. Will., Proceedings of the Royal Society of Victoria ser. 2,
33: 146, pl. VI (1921); Pultenaea tenuifolia var. recurvifolia Benth., Fl. Austral. 2: 140 (1864).
Type citation: ‘Near Portland, Allitt.’
Type: Cape Nelson, near Portland, Victoria, W. Allitt s.n. (lecto, here designated: MEL
2057338; isolecto: BM 000544779, K 000118046, K 000118047, NSW 38979).
Typification: Bentham had two collections available on loan from MEL, but only cited one
collection (Near Portland, Allitt) [
62
], with two sheets now located at K and two other sheets
at BM and MEL. Only the sheet at MEL bears the name ‘var. recurvifolia’, but this does not
appear to be in Bentham’s script, though, unfortunately, the bottom of the label has been
cut off at some point. The sheets at BM and K bear the name P. recurvifolia, indicating that
they were distributed much later. For this reason, we choose the MEL sheet as lectotype. A
second collection (Mouth of the Glenelg [River], W. Allitt s.n. (K 000118045)) was annotated
as the ‘holotype’ by M.D. Crisp in 1982, but it was not cited by Bentham [
62
], so it has no
type status.
Notes: de Kok & West [
94
] erroneously treated this taxon as a synonym of P. daltonii, a later
name. Further investigation has shown that P. recurvifolia is more closely allied to P. hispidula
R.Br. ex Benth., as noted in VicFlora ([
133
], updated by Stajsic 2019). Its taxonomic status
requires further investigation, and it is uncertain whether the taxon should be reinstated or
included as a synonym of P. hispidula.
Stonesiella
Crisp & P.H. Weston in M.D. Crisp et al., Taxon 48(4): 711 (1999). Type:Stonesiella
selaginoides (Hook.f.) Crisp & P.H. Weston
Urodon
Turcz., Bulletin de la Societe Imperiale des Naturalistes de Moscou 22(3): 16 (1849). Type:
Urodon capitatus Turcz.
Mirbelieae s.l.
Daviesia
Sm., Transactions of the Linnean Society of London 4: 220 (1798). Type:Daviesia
acicularis Sm., designated by J. Hutchinson, Dicotyledones.The Genera of Flowering Plants
(Angiospermae) 1: 339 (1964).
Erichsenia
Hemsl., Hooker’s Icones Plantarum 28: t. 2777 (1905). Type:Erichsenia uncinata
Hemsl.
Gompholobium
Sm., Transactions of the Linnean Society of London 4: 220 (1798). Type:Gom-
pholobium grandiflorum Sm., designated by J.A. Chappill et al., Australian Systematic Botany
21(2): 68 (2008).
Burtonia R.Br. in W.T. Aiton, Hortus Kewensis Edn. 2, 3: 12 (1811), nom. cons., non Salisb.
(1807), nom. rej.Weihea Rchb., Conspectus Regni Vegetabilis 212
b
(1828), nom. illeg., non
Sprengel (1825), nom. rej.Type:Burtonia scabra (Smith) W.T. Aiton
Gompholobium sect. Disporaea F. Muell., Fragmenta Phytographiae Australiae 3(18): 29 (1862),
nom. inval., nom. nud.
Isotropis
Benth. in S.F.L. Endlicher et al., Leguminosae. Enumeratio plantarum quas in Novae
Hollandiae ora austro-occidentali ad fluvium Cygnorum et in Sinu Regis Georgii collegit Carolus
liber baro de Hügel 28 (1837). Type:Isotropis striata Benth. [= I. cuneifolia (Sm.) Walp.].
Notes: Bentham [
59
] described two species in his new genus Isotropis,I. biloba Benth. and I.
striata Benth. but did not designate either as the type as this was not a practice at the time.

Diversity 2021,13, 391 32 of 38
Bentham [
62
] recognised I. striata as a good species, but included I. biloba as a synonym,
having examined a broader range of specimens since his original treatment. On that basis,
we here designate I. striata as the type of the genus.
The application of both names has been considered uncertain in Australian literature,
but one of us (MDC) examined the type material of each taxon at W in 1982, and these
specimens have recently become available online. They are both confirmed as synonyms of
I. cuneifolia, as indicated by Bentham ([
62
]; as Callistachys cuneifolia Sm.). Both names are
applicable to I. cuneata subsp. cuneata based on the characters defined by Keighery [
113
],
though this taxon is variable and additional taxa are likely to be recognised.
Sphaerolobium
Sm. in K.D.E. Koenig, & J. Sims (eds), Annals of Botany 1(3): 509 (1805).
Type:Sphaerolobium vimineum Sm.
Roea Hügel ex Benth. in S.F.L. Endlicher et al., Leguminosae. Enumeratio plantarum quas in
Novae Hollandiae ora austro-occidentali ad fluvium Cygnorum et in Sinu Regis Georgii collegit
Carolus liber baro de Hügel 34 (1837). Sphaerolobium sect. Roea (Hügel ex Benth.) Benth.,
Flora Australiensis 2: 63, 64 (1864). Type:Roea linophylla Hügel ex Benth. [= Sphaerolobium
linophyllum (Hügel ex Benth.) Benth.].
Viminaria
Sm. in K.D.E. Koenig, & J. Sims (eds), Annals of Botany 1(3): 507 (1805). Type:
Viminaria denudata Sm., nom. illeg. [= V. juncea (Schrad. & J.C. Wendl.) Hoffmanns.].
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