Understanding Diversity and Systematics in Australian Fabaceae Tribe Mirbelieae

Peter Jobson

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 ...

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 1 and Peter H. Weston 1 1 2 3 4 5 * 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 National 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.) School of Biological Sciences, The University of Queensland, Brisbane, QLD 4072, Australia; l.cook@uq.edu.au Research School of Biology, The Australian National University, Acton, ACT 2601, Australia; Mike.Crisp@anu.edu.au Northern 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 Australian 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; typification; new combinations published maps and institutional affiliations. 1. Introduction 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/). 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 Southwest 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 megametophytes 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); embryosac 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] Jacksonia Leptosema Latrobea Chorizema Oxylobium Mirbelia Callistachys Aotus Otion Urodon Phyllota Euchilopsis Dillwynia Eutaxia Almaleea Stonesiella Pultenaea Podolobium Gastrolobium Isotropis Mirbelieae “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ Mirbelia group “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ Isotropis Erichsenia “ Daviesia group Viminaria Daviesia Sphaerolobium Gompholobium Bossiaea Platylobium Goodia Aenictophyton Muelleranthus Ptychosema Paragoodia “ “ “ “ Bossiaeeae “ “ “ “ “ “ “ “ “ “ Bossiaeeae “ “ “ “ “ “ 5-nucleate “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ giant antipodals “ “ “ “ “ “ “ “ “ “ “ Current Paper Monophyletic cpDNA Monophyletic ITS Core Mirbelieae “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ “ Mirbelieae s.l. yes yes yes no no no – no – – yes – no yes – – no no yes * yes yes yes yes no no no – no – – yes – no no – – no no yes * yes “ – – “ “ “ “ Bossiaeeae “ “ “ “ “ “ – yes yes no no yes yes – – – – – yes yes no no yes yes – – – – *: 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], Gastrolobium 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 revisions 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 complexes 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 A of 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 Callistachys 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 Australian 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 understanding 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, diversification 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 (A) Figure 1. Cont. Diversity 2021, 13, 391 7 of 38 (B) Figure 1. Cont. Diversity 2021, 13, 391 8 of 38 (C) Figure 1. Cont. Diversity 2021, 13, 391 9 of 38 (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. 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 Almaleea Crisp & P.H.Weston A. cambagei (Maiden & E.Betche) Crisp & P.H.Weston Aotus Sm. A. carinata Meisn. A. cordifolia Benth. A. sp. Esperance (P.G. Wilson 7904) A. subglauca Blakey & McKie Callistachys Vent. C. lanceolata Vent. Chorizema Labill. C. aciculare (DC.) C.A.Gardner C. carinatum (Meisn.) J.M.Taylor & Crisp C. dicksonii Graham C. genistoides (Meisn.) C.A.Gardner C. obtusifolium (Sweet) J.M.Taylor & Crisp C. parviflorum Benth. C. rhombeum R.Br. C. ulotropis J.M.Taylor & Crisp C. varium Benth. Dillwynia Sm. D. parvifolia R.Br. ex Sims D. phylicoides A.Cunn. Euchilopsis F.Muell. E. linearis (Benth.) F.Muell. Eutaxia R.Br. Collector Number Herbarium GenBank M.D. Crisp 9197 CANB AF113775 J.A. Chappill J.A. Chappill M.D. Crisp M.D. Crisp 6581 6587 9197 9047 PERTH PERTH CANB CANB AY883181 AY883182 AF518163 AF113776 G.T. Chandler 474 CANB AY015072 M.D. Crisp M.D. Crisp M.D. Crisp M.D. Crisp M.D. Crisp M.D. Crisp M.D. Crisp M.D. Crisp M.D. Crisp 9202 9237 9166 9026 9201 9166 9230 9214 8528 CANB CANB CANB CANB CANB CANB CANB CANB CANB AF518149 AF518150 AY251250 AF518151 AY251251 AF518152 AF518153 AY251253 AF518154 J.M. Taylor M.D. Crisp 360 9049 CBG CANB AF113777 AF113778 M.D. Crisp 8535 CANB AF113779 Diversity 2021, 13, 391 11 of 38 Table 2. Cont. Taxon Collector Number Herbarium GenBank C. racemosum (Meisn.) J.M.Taylor & Crisp E. microphylla (R.Br.) C.H.Wright & Dewar E. neurocalvx (Turcz.) Chappill & G.R.Hend. Gastrolobium R.Br. G. alternifolium G.Chandler & Crisp G. bilobum R.Br. G. bracteolosum (F.Muell.) G.Chandler & Crisp G. brownii Meisn. G. calycinum Benth. G. celsianum (Lem.) G.Chandler & Crisp G. congestum G.Chandler & Crisp G. coriaceum (Sm.) G.Chandler & Crisp G. cuneatum Henfr. G. ebracteolatum G.Chandler & Crisp G. formosum (Kippist ex Lindl.) G.Chandler & Crisp G. grandiflorum F.Muell. G. hookeri Meisn. G. latifolium (R.Br.) G.Chandler & Crisp G. leakeanum J.Drumm. G. luteifolium (Domin) G.Chandler & Crisp G. melanopetalum (F.Muell.) G.Chandler & Crisp G. minus (Crisp) G.Chandler & Crisp G. modestum (Crisp) G.Chandler & Crisp G. obovatum Benth. G. parviflorum (Benth. ex Lindl.) Crisp G. plicatum Turcz. 1 G. plicatum Turcz. 2 G. praemorsum (Meisn.) G.Chandler & Crisp G. pulchellum Turcz. G. pusillum Crisp & P.H.Weston G. pyramidale T.Moore G. reticulatum (Meisn.) Benth. G. sericeum (Sm.) G.Chandler & Crisp G. spathulatum Benth. ex Lindl. G. spinosum Benth. ex Lindl. G. subcordatum (Benth.) G.Chandler & Crisp G. truncatum Benth. G. vestitum (Domin) G.Chandler & Crisp Jacksonia Sm. J. alata Benth. J. horrida DC. J. macrocalyx Meisn. Latrobea Meisn. L. brunonis (Benth.) Meisn. L. genistoides (Meisn.) Meisn. L. hirtella (Turcz.) Benth. Leptosema Benth. L. aphyllum (Hook.) Crisp L. daviesioides (Turcz.) Crisp Mirbelia Sm. M. baueri (Benth.) Joy Thomps. M. confertiflora Pedley M. depressa E.Pritz. M. dilatata R.Br. M. longifolia C.A.Gardner 1 M.D. Crisp M.D. Crisp M.D. Crisp 9012 8918 8525 CANB CANB CANB AY251252 AF113780 AF113789 M.D. Crisp G.T. Chandler G.T. Chandler G.T. Chandler G.T. Chandler M.D. Crisp G.T. Chandler G.T. Chandler M.D. Crisp M.D. Crisp 8512 724 426 726 544 9009 404 723 8937 8471 CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB AY015088 AY015073 AY015063 AY015074 AY015075 AY015064 AY015076 AY015089 AY015077 AY015102 M.D. Crisp 8933 CANB AY015085 G.T. Chandler M.D. Crisp G.T. Chandler M.D. Crisp M.D. Crisp M.D. Crisp M.D. Crisp M.D. Crisp G.T. Chandler G.T. Chandler G.T. Chandler M.D. Crisp G.T. Chandler M.D. Crisp M.D. Crisp G.T. Chandler G.T. Chandler J.M. Taylor M.D. Crisp G.T. Chandler M.D. Crisp M.D. Crisp M.D. Crisp 598 8907 365 8481 9407 8470 8922 8465 657 760 623 9014 729 8480 8921 488 540 1959 8448 548 8511 8919 8489 CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CBG CANB CANB CANB CANB CANB AY015078 AY015090 AY015065 AY015091 AY015092 AY015066 AY015067 AY015068 AY015093 AY015079 AY015094 AF518161 AY015069 AY015095 AY015080 AY015096 AY015097 AY015070 AY015098 AY015081 AY015071 AY015082 AY015099 M.D. Crisp M.D. Crisp M.D. Crisp 8956 8934 9272 CANB CANB CANB AF518146 AY015084 AF518147 J.A. Chappill J.A. Chappill M.D. Crisp 6564 6567 8478 PERTH PERTH CANB AY883186 AY883187 AF113781 M.D. Crisp M.D. Crisp 9019 9193 CANB CANB AF518148 AY883188 M.D. Crisp M.D. Crisp M.D. Crisp M.D. Crisp M.D. Crisp 9144 9050 9020 8491 9263 CANB CANB CANB CANB CANB AY251254 AF518155 AY015086 AY015087 AY883189 Diversity 2021, 13, 391 12 of 38 Table 2. Cont. Taxon Collector Number Herbarium GenBank M. longifolia C.A.Gardner 2 M. microphylla (Turcz.) Benth. M. oxylobioides F.Muell. 1 M. oxylobioides F.Muell. 2 M. oxylobioides F.Muell. 3 M. pungens A.Cunn. ex G.Don. M. ramulosa (Benth.) C.A.Gardner M. rhagodioides Crisp & J.M.Taylor M. rubiifolia (Andrews) G.Don M. seorsifolia (F.Muell.) C.A.Gardner M. sp. Pedicellosa (A.Monro 25) M. sp. Ternata (M.D.Crisp & L.G.Cook MDC 9267) M. speciosa Sieber ex DC. subsp. speciosa 1 M. speciosa Sieber ex DC. subsp. speciosa 2 M. viminalis (A.Cunn. ex Benth.) C.A.Gardner ‘Otion’ Crisp & P.H.Weston ined. ‘O. microphyllum’ (Benth.) Crisp & P.H.Weston ined. Oxylobium Andr. O. arborescens R.Br. 1 O. arborescens R.Br. 2 O. cordifolium Andrews O. ellipticum (Vent.) R.Br. 1 O. ellipticum (Vent.) R.Br. 2 O. lineare (Benth.) Benth. O. pulteneae DC. O. robustum Joy Thomps. Phyllota (DC.) Benth. P. phylicoides (Sieber ex DC.) Benth. 1 P. phylicoides (Sieber ex DC.) Benth. 2 Podolobium R.Br. P. aciculiferum F.Muell. P. aestivum Crisp & P.H.Weston P. alpestre (F.Muell.) Crisp & P.H.Weston P. ilicifolium (Andrews) Crisp & P.H.Weston P. procumbens (F.Muell.) Crisp & P.H.Weston P. scandens (Sm.) DC. Pultenaea Sm. P. adunca Turcz. P. alea de Kok P. arida E.Pritz. 1 P. arida E.Pritz. 2 P. aspalathoides Meisn. P. baeuerlenii F.Muell. P. barbata C.R.P.Andrews P. phylicoides (Sieber ex DC.) Benth. 1 P. phylicoides (Sieber ex DC.) Benth. 2 Podolobium R.Br. P. aciculiferum F.Muell. P. aestivum Crisp & P.H.Weston P. alpestre (F.Muell.) Crisp & P.H.Weston P. ilicifolium (Andrews) Crisp & P.H.Weston P. procumbens (F.Muell.) Crisp & P.H.Weston P. scandens (Sm.) DC. Pultenaea Sm. P. adunca Turcz. P. alea de Kok M.D. Crisp A. Monro M.D. Crisp Yi Yi M.D. Crisp A. Monro M.D. Crisp ANBG H. King A. Monro 9263 22 9112 14,825 14,825 9138 4 9259 8406509 314 25 CANB CANB CANB KUN KUN CANB CANB CANB CANB CANB CANB AY251255 AF518156 AY251264 NC_047371 MN709855 AY251257 AY251258 AY251259 AF518157 AY251260 AY251256 M.D. Crisp 9266 CANB AY251262 M.D. Crisp – D. Morris 9123 8100876 547 CANB CBG CANB AY251261 AF518158 AY251263 M.D. Crisp 8970 CANB AF113782 G.T. Chandler M.D. Crisp M.D. Crisp M.D. Crisp G.T. Chandler M.D. Crisp M.D. Crisp I.R. Telford 616 9093 9133 9092 603 8471 9046 4294 CANB CANB CANB CANB CANB CANB CANB CBG AY015100 AF113783 AF518159 AF113784 AY015101 AY015102 AY015103 AY015104 M.D. Crisp M.R. Gillings 9048 Soil core CANB – AF113785 JN392738 G.T. Chandler G.T. Chandler G.T. Chandler G.T. Chandler B. Hadlow G.T. Chandler 606 612 1039 308 461 309 CANB CANB CANB CANB CBG CANB AF518160 AY015105 AY015106 AY015107 AY015108 AY015109 J.A. Chappill J. Westaway J.A. Chappill L.A. Orthia L.A. Orthia B. Pfeil T.R. Lally M.D. Crisp M.R. Gillings 6544 924 6272 71 50 263 1309 9048 Soil core PERTH CANB PERTH CANB CANB CANB CANB CANB – AY883190 AY883191 AF518162 AY883192 AY883193 AY883194 AY883195 AF113785 JN392738 G.T. Chandler G.T. Chandler G.T. Chandler G.T. Chandler B. Hadlow G.T. Chandler 606 612 1039 308 461 309 CANB CANB CANB CANB CBG CANB AF518160 AY015105 AY015106 AY015107 AY015108 AY015109 J.A. Chappill J. Westaway 6544 924 PERTH CANB AY883190 AY883191 Diversity 2021, 13, 391 13 of 38 Table 2. Cont. Taxon Collector Number Herbarium GenBank P. arida E.Pritz. 1 P. arida E.Pritz. 2 P. aspalathoides Meisn. P. baeuerlenii F.Muell. P. barbata C.R.P.Andrews P. blakelyi Joy Thomps. P. brachyphylla Turcz. P. brachytropis Benth. P. calycina (Turcz.) Benth. subsp. calycina P. calycina subsp. proxena Orthia & Chappill P. costata H.B.Will. P. daena Orthia P. daphnoides J.C.Wendl. P. densifolia F.Muell. P. dentata Labill. P. divaricata H.B.Will. P. echinula Sieber ex DC. P. elachista (F.Muell.) Crisp P. empetrifolia Meisn. 1 P. empetrifolia Meisn. 2 P. empetrifolia Meisn. 3 P. empetrifolia Meisn. 4 P. ericifolia Benth. 1 P. ericifolia Benth. 2 P. ericifolia Benth. 3 P. ericifolia Benth. 4 P. ericifolia Benth. 5 P. ericifolia Benth. 6 P. euchila DC. P. fasciculata Benth. P. flexilis Sm. P. glabra Benth. P. graveolens Tate P. gunnii Benth. P. heterochila F.Muell. 1 P. heterochila F.Muell. 2 P. hispidula Benth. P. indira Orthia & Crisp subsp. indira 1 P. indira Orthia & Crisp subsp. indira 2 P. indira Orthia & Crisp subsp. indira 3 P. indira subsp. monstrosita Orthia P. juniperina Labill. P. kraehenbuehlii P.J.Lang P. largiflorens F.Muell. ex Benth. P. laxiflora Benth. P. luehmannii Maiden P. muelleri Benth. P. myrtoides A.Cunn. P. ochreata Meisn. P. parviflora Sieber ex DC. P. patellifolia H.B.Will. P. pauciflora M.B.Scott 1 P. pauciflora M.B.Scott 2 P. pauciflora M.B.Scott 3 P. pedunculata Hook. P. petiolaris A.Cunn. ex Benth. P. pinifolia Meisn. P. purpurea (Turcz.) Crisp & Orthia J.A. Chappill L.A. Orthia L.A. Orthia B. Pfeil T.R. Lally R.P.J. de Kok L.A. Orthia L.A. Orthia L.A. Orthia L.A. Orthia R.P.J. de Kok L.A. Orthia E. Gauba J. Mant M.D. Crisp F.E. Davies R.P.J. de Kok L.A. Orthia L.A. Orthia L.A. Orthia L.A. Orthia L.A. Orthia L.A. Orthia L.A. Orthia L.A. Orthia L.A. Orthia L.A. Orthia M.D. Crisp D.A. Taylor R.P.J. de Kok R.P.J. de Kok R.P.J. de Kok R.P.J. de Kok R.P.J. de Kok J.A. Chappill D.A. Taylor R.P.J. de Kok L.A. Orthia L.A. Orthia M.D. Crisp L.A. Orthia R.P.J. de Kok R.J. Bayer D.L.Jones A. Monro R.P.J. de Kok R.P.J. de Kok D.A. Taylor L.A. Orthia R.P.J. de Kok R.P.J. de Kok L.A. Orthia L.A. Orthia L.A. Orthia R.P.J. de Kok R.P.J. de Kok L.A. Orthia L.A. Orthia 6272 71 50 263 1309 715 56 37 51 63 730 57 22,264 71 9053 1887 709 66 54 53 79 80 88 39 41 46 48 8451 55 860 705 cultivated 778 900 4279 1569 780 59 70 9178 78 713 SA99010 15,757 64 738 779 7 42 793 761 83 84 85 756 903 40 60 PERTH CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CBG CANB CANB CBG CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB – CANB CANB PERTH CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB AF518162 AY883192 AY883193 AY883194 AY883195 AY883196 AY883197 AY883198 AY883199 AY883199 AY883200 AY883201 AF113786 AY883202 AY015110 AY883203 AY883204 AY883205 AY883206 AY883385 AY883386 AY883387 AY883207 AY883388 AY883389 AY883390 AY883391 AF113788 AY883208 AY883209 AY883210 AY883211 AY883212 AY883213 AY883231 AY883401 AY883214 AY883215 AY883395 AY883396 AY883397 AY883216 AY883217 AY883218 AY883219 AY883220 AY883221 AY883222 AY883223 AY883224 AY883225 AY883400 AY883226 AY883399 AY883227 AY883228 AY883229 AY883230 Diversity 2021, 13, 391 14 of 38 Table 2. Cont. Taxon Collector Number Herbarium GenBank P. radiata H.B.Will. P. reticulata (Sm.) Benth. P. rosmarinifolia Lindl. P. rotundifolia (Turcz.) Benth. P. scabra R.Br. P. sericea (Benth.) Corrick P. setulosa Benth. P. skinneri F.Muell. P. sp. Mt Lesueur (Beard 7827) P. spinosa (DC.) H.B.Will. P. spinulosa (Turcz.) Benth. P. stipularis Sm. P. stricta Sims P. strobilifera Meisn. 1 P. strobilifera Meisn.2 P. strobilifera Meisn.3 P. subalpina (F.Muell.) Druce P. subspicata Benth. P. tarik de Kok P. tenuifolia R.Br. & Sims 1 P. tenuifolia R.Br. & Sims 2 P. trichophylla H.B.Will. ex J.M.Black P. trifida J.M.Black P. trinervis J.M.Black P. tuberculata Pers. P. verruculosa Turcz. P. vestita R.Br. ex Aiton P. victoriensis Corrick P. villifera Sieber ex. DC. P. viscidula Tate P. vrolandii Maiden P. whiteana S.T.Blake P. williamsoniana J.H.Willis P. wudjariensis Orthia Stonesiella Crisp & P.H.Weston S. selaginoides (Hook.f.) Crisp & P.H.Weston Urodon Turcz. U. capitatus Turcz. OUTGROUPS Daviesia Sm. D. elliptica Crisp D. flava Pedley D. longifolia Benth. Sphaerolobium Sm. S. medium R.Br. S. minus Labill. S. nudiflorum (Meisn.) Benth. L.A. Orthia L.A. Orthia D.A. Taylor L.A. Orthia R.P.J. de Kok J. Mant R.P.J. de Kok L.A. Orthia J.S. Beard R.P.J. de Kok L.A. Orthia R.P.J. de Kok R.P.J. de Kok L.A. Orthia L.A. Orthia L.A. Orthia R.P.J. de Kok R.P.J. de Kok R.P.J. de Kok L.A. Orthia R.P.J. de Kok R.P.J. de Kok R.P.J. de Kok R.P.J. de Kok R.P.J. de Kok L.A. Orthia R.P.J. de Kok R.P.J. de Kok R.P.J. de Kok R.P.J. de Kok R.P.J. de Kok M.D. Crisp R.P.J. de Kok C.E. Woolcock 38 47 68 61 909 72 716 36 7827 cultivated 68 701 729 44 52 75 721 718 666 64 803 838 819 801 702 45 830 762a 828 833 784 9113 754 2250 CANB CANB CANB CANB CANB CANB CANB CANB PERTH – CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CANB CBG AY883232 AY883233 AY883234 AY883235 AY883236 AY883237 AY883238 AY883239 AY883398 AY883240 AY883241 AY883242 AY883243 AY883392 AY883393 AY883394 AY883244 AY883245 AY883246 AY883247 AY883248 AY883249 AY883250 AY883251 AY883252 AY883253 AY883254 AY883255 AY883256 AY883257 AY883258 AY883259 AY883260 AY883261 R. Burns 258 CBG AF113791 M.D. Crisp 8523 CANB AF113792 M.D. Crisp I.R.D. Telford M.D. Crisp 9051 12,054 9246 CANB CBG CANB AF518130 KY426177 KY426196 M.D. Crisp M.D. Crisp R. Butcher 8942 9054 1229A CANB CANB PERTH AF518136 AF518135 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 Sequence length range (bp) Aligned length (bp) GC content (%) Variable sites Gaps + missing characters (%) 200 518–934 934 32.2 680 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. Significantly, 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 hybridisation, 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 Mirbelieae 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) Core Mirbelieae Almaleea * Aotus s.s. * Aotus Clade A * Aotus Clade B * Aotus Clade C Callistachys s.l. * Chorizema s.l. * Dillwynia Euchilopsis * Eutaxia s.s. Gastrolobium Jacksonia * Latrobea Leptosema * Mirbelia * ’Otion’ ined. Oxylobium * Phyllota * Podolobium Pultenaea s.s. * Pultenaea Clade A * Pultenaea Clade B * Pultenaea Clade C * Pultenaea Clade D * Sclerothamnus R.Br. Stonesiella Urodon Mirbelieae s.l. Daviesia Erichsenia Gompholobium Isotropis Sphaerolobium Viminaria Total Mirbelieae s.l. Number of Species No. of Species with DNA Sequence Data/No. of Sequences Published Descriptions Available New or Revised Descriptions Required 5 6 3 3 10 5 28 36 1 12 109 74 9 12 26 5 7 10 3 130 1 2 9 6 12 1 4 1/4 4/6 2/5 0 0 4/17 18/32 3/7 1/4 3/6 82/313 6/15 4/12 2/5 15/65 1/2 6/40 2/7 3/17 58/222 1/3 2/6 9/28 6/18 0 1/1 2/4 5 0 1 2 0 3 28 4 0 12 109 74 3 12 28 1 1 1 3 83 1 2 7 6 10 1 0 0 6 2 1 10 2 0 32 1 0 0 0 6 0 8 4 6 9 0 47 0 0 2 0 2 0 4 130 1 46 15 22 1 130/419 1/3 5/9 4/18 3/7 1/4 130 0 46 3 22 1 0 1 0 12 0 0 755 385/1298 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 circumscriptions. 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 diversity 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 maintained 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 Figure 2. Morphological diversity of representative species from Fabaceae tribe Bossiaeeae (sister to Mirbelieae) and non 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 Figure 3. Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae). (A) Almaleea 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 Figure 4. Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae). (A 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)). Diversity 2021, 13, 391 23 of 38 Figure 5. Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae). (A) Jacksonia 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 Figure 6. Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae). (A) OxyFigure 6. Morphological diversity of representative species from Fabaceae tribe Mirbelieae (core Mirbelieae). (A) Oxylobium 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)). 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. Numerous 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 syntypes. 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: Orthotropis 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: Gompholobium 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 212b (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.) 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