The Callerya Group redefined and Tribe Wisterieae (Fabaceae) emended based on morphology and data from nuclear and chloroplast DNA sequences

Abstract The Tribe Wisterieae (Zhu 1994), founded on the single genus Wisteria, is emended and recircumscribed based on morphology and data from nuclear ITS and ndhJ-trnF, matK and rbcL chloroplast DNA sequences. This newly enlarged tribe comprises 36 species and 9 infraspecific taxa within 13 described genera. Six genera are new, two are reinstated and five were previously placed in Tribe Millettieae. The genus Adinobotrys is also reinstated comprising two species including the new combination A.vastus. Other reinstated genera include Whitfordiodendron, with four species, and Padbruggea, with three species, including the reinstatement of P.filipes and the new combination P.filipesvar.tomentosa. The existing genera Afgekia, Callerya, Endosamara (with the new combination E.racemosavar.pallida), Sarcodum and Wisteria, with the new combinations W.frutescenssubsp.macrostachya are evaluated. The new genera comprise three Australasian species in Austrocallerya: A.australis, A.megasperma and A.pilipes; Wisteriopsis with five species from east Asia has six new combinations: W.japonica, W.kiangsiensis, W.championii, W.eurybotrya, W.reticulata and W.reticulatavar.stenophylla. Two species comprise the new Thai genus Kanburia: K.tenasserimensis and K.chlorantha. Nanhaia comprises the two species: N.fordii and N.speciosa and the monotypic genera Sigmoidala and Serawaia are based respectively on the species S.kityana and S.strobilifera. Lectotypes are designated for the names Adinobotrysfilipes, A.myrianthus, Millettiabonatiana, Millettiabracteosa, Millettiachampionii, Millettiacinerea, Millettiadielsiana, Millettiakityana, M.maingayi, Millettianitida, Millettiaoocarpa, Millettiapurpurea, M.reticulata, M.reticulatavar.stenophylla, Padbruggeadasyphylla, Pterocarpusaustralis, Robiniaracemosa, Whitfordiodendronscandens, W.sumatranum and Wisteriapallida. A neotype is designated for the name Millettialeiogyna.


Introduction
The Tribe Millettieae was first described by Miquel (1855: 137), based on the type genus Millettia Wight & Arn. (Wight and Arnott 1834: 263). This genus of six species was characterised largely by the pods of the southern Indian type species M. rubiginosa Wight & Arn. Miquel emphasised the compressed nature of the pods as a significant distinguishing character and his tribal description very loosely defined the new Tribe Millettieae, which included eight genera: Brachypterum (Wight & Arn.) Benth., Derris Lour., Pongamia Adans., Padbruggea Miq., Aganope Miq., Millettia, Otosema Benth. and Mundulea Benth. (Miquel 1855: 137). Miquel (1855: 137)  Millettia was, furthermore, distinguished from the genera Pongamia Adans. and Dalbergia L.f. by the legume being compressed around the seeds and by the fruit's thick woody texture (Wight and Arnott 1834: 263). Dunn (1912a) in his revision of Millettia placed the genus in Tribe Galegeae (Bronn) Torr. & Gray subtribe Tephrosiinae Benth. (as Subtribe "Tephrosieae"). Geesink (1984: 3) described Tribe Millettieae with the characters: Inflorescence of panicles, pseudopanicles or derived pseudopanicles; wing petals adherent to the keel; keel petals usually valvately connate; pod dehiscent or indehiscent; seed chamber mostly absent; seeds 1 or few; and without any uniquely defining character". Geesink (1984: 4) admitted that "the contents of this chapter [Delimitations of Millettieae and related tribes] will be disappointing for those who expect a final answer to the questions suggested by the title".
Geesink's major revision of this alliance, which he much enlarged to comprise 43 genera within Tribe Millettieae, was clearly polyphyletic with an assemblage of taxa having a range of unifying as well as contradicting characters. Although far from definitive, this revision was a major step forward and did lay a sound basis for subsequent research in the tribe. Geesink's (1984) generic treatment included a set of genera in the "Callerya Group" (Hu et al. 2002;Hu and Chang 2003) : Wisteria Nutt. (1818), Callerya Endl. (1843), Afgekia Craib (1927) and Endosamara Geesink (1984). Subsequently, Sarcodum Lour. (1790) was also placed in this group (Schrire 2005;Clark 2008) (see Table 1). The genus Antheroporum Gagnep., placed tentatively in this grouping by Schrire (2005), has subsequently been shown to belong within the core-Millettieae (LPWG 2016;Mattapha 2017: 53).
Over the past 30 years a large number of DNA-based phylogenies have analysed many taxa from Tribe Millettieae (Palmer et al. 1987;Lavin et al. 1990;Doyle et al. 1997;Lavin et al. 1998;Doyle et al. 2000;Kajita et al. 2001;Hu et al. 2000;Hu et al. 2002;Hu and Chang 2003;Wojciechowski et al. 2004;Schrire 2005;Schrire et al. 2009;Wink 2013;Li et al. 2014;de Quieroz et al. 2015). Analysis of data from the phytochrome gene family PHY (Lavin et al. 1998) has shown that a core-Millettieae group is monophyletic and may be defined by the presence of pseudoracemes and pseudopanicles. Moreover, while the millettioid-phaseoloid alliance as a whole falls within the large non-protein amino acid accumulating (NPAAA) clade (Wojciechowski et al. 2004;Cardoso et al. 2012;Wink 2013;Wojciechowski 2013;De Quieroz et al. 2015), Lavin et al. (1998) showed that the core-Millettieae group are diagnosed by a loss of the ability to accumulate the non-protein amino acid canavanine. In these studies the Callerya group does not belong with the Millettioid group but rather is accommodated in the Hologalegina clade . Lavin et al. (1998) also revealed that Afgekia, Callerya, Endosamara and Wisteria, i.e. a significant part of the Callerya group, did accumulate canavanine rather than alkaloids in their seeds and that they all possessed either true panicles or true racemes. Furthermore, Lavin et al. (1998) postulated that owing to the presence of true racemes, Sarcodum was also likely to accumulate canavanine and would therefore not be part of the core-Millettioid group. The morphological distinction between true and pseudoracemes is that in "true" racemes the flowers are inserted singly on the rachis (the unit comprising a flower, pedicel and bract). Pseudoracemes (Lackey 1981) on the other hand, consist of more than one flower inserted at a node on the rachis (the unit comprising two or more flowers, pedicels and bracts all subtended by a secondary bract representing branch reduction). Racemes and pseudoracemes are further compounded into panicles and pseudopanicles.
The Callerya group occurs in a more inclusive subset of taxa that all lack one copy of chloroplast DNA, the Inverted Repeat Lacking Clade or IRLC (Palmer et al. 1987;Lavin et al. 1990;Liston 1995;Doyle et al. 1997;Wojciechowski et al. 2000). The loss of a prominent inverted repeat structure in cpDNA in legumes had previously been observed in the genera Vicia (Koller and Delius 1980) and Pisum (Palmer and Thompson 1981). The genome of Wisteria was also discovered to have deleted one half of the inverted repeat amounting to 25 kb of DNA (Palmer et al. 1987). It was apparent that whereas the other legume genera were rearranged genetically as a result of the loss of the inverted repeat, both Wisteria floribunda and Medicago sativa remained otherwise unrearranged (Palmer et al. 1987). The IRLC is sister to Tribes Loteae, Sesbanieae and Robinieae (Lewis et al. 2005), which retain the inverted repeat Cardoso et al. 2012;Cardoso et al. 2013;LPWG 2013LPWG , 2017. Additional evidence from chloroplast rbcL sequence data has also revealed that the Millettieae lie outside the IRLC (Lavin et al. 1990;Doyle et al. 1997;Kajita et al. 2001;Hu and Chang 2003), while the Callerya group all fall within the IRLC. These data refute the previously made assumptions that the group belongs with the Millettieae. Lavin et al. (1990) noted that although Wisteria and Millettia japonica both showed hypogeal seed germination and a lianescent habit, which are characteristic of many Millettieae genera, these species differed from the Millettieae in their wholly temperate distribution, the lack of the inverted repeat and both had a base chromosome number of x = 8 as opposed to x = 11 or 12. The analyses of Hu and Chang (2003), based on plastid rbcL sequence data, confirmed that Afgekia sericea, Callerya vasta, Endosamara racemosa, Millettia japonica and two Wisteria species all belonged within the large IRLC. Their results, however, were based on comparatively limited taxon sampling of taxa within the Callerya group. Wink (2013) examined 1276 species of Leguminosae for the distribution of secondary metabolites mapped against phylogenetic trees generated by combined sequence data from cpDNA rbcL, matK and nrDNA ITS. In the study, it was shown that Wisteria and Callerya nested within the IRLC and that they possessed isoflavones in common with most but not all other taxa within the IRLC.
A unique marker further distinguishes the Callerya group, adding weight to the distinctiveness of this assemblage of genera. Jansen et al. (2008) undertook a comprehensive survey for the retention or loss of two chloroplast introns among 301 legume species representing three subfamilies and 198 genera. Their survey of the presence or absence of the rps12 intron revealed that along with 49 of the millettioid-phaseoloids sampled from outside the IRLC, Afgekia filipes, A. sericea, Callerya atropurpurea, C. australis, C. megasperma, C. pilipes, Endosamara racemosa, Millettia (sic) japonica, Wisteria brachybotrys, W. floribunda, W. frutescens, W. macrostachya and W. sinensis -each from inside the IRLC -all retained the intron. Of the 77 other taxa sampled from within the IRLC all -without exception -had lost the intron. Significantly, therefore, all genera within the IRLC surveyed for the presence or absence of the rps12 intron showed it to be lacking, except for the Callerya group, marking out the latter as unique within the IRLC (Jansen et al. 2008). Seven species of Glycyrrhiza surveyed by Jansen et al. (2008), whose position in recent phylogenies (Doyle et al. 2000;Lewis et al. 2005;LPWG 2013LPWG , 2017Li et al. 2014), was placed sister to the Callerya group within the IRLC, all lacked the rps12 intron. Glycyrrhiza, which is represented in our analyses, has therefore not been included as part of the Callerya group.
The Callerya group is thus uniquely diagnosed by a combination of lacking the 25 kb. inverted repeat of cpDNA and possessing the cpDNA rps12 intron. Representa- Table 1.
Published treatments of species in the Callerya group based on their assignment to genus, from 1984 to the present. Those taxa highlighted in bold represent Chinese species of Callerya s.l. for which we were unable to see material. All species epithets are transferable across genera for comparison purposes. tives of Afgekia, Callerya and Wisteria from this subgroup of taxa have also been found to group together according to data from sequences of nuclear DNA ITS spacer regions (Hu et al. 2002;Li et al. 2014). Zhu (1994) defined her new Tribe Wisterieae comparing only pollen from four species of Wisteria to the millettioid-phaseoloid genera Craspedolobium Harms, Derris, Millettia, Pongamia and Tephrosia Pers. The genus Pongamia Adans. is now considered synonymous with Millettia Wight & Arn. (Schrire 2005: 383). The pollen grains of the four Wisteria species exhibited much broader polar regions (apocolpia) and a distinctive reticulate pollen surface compared to the other taxa examined. Zhu (1994) also made comparisons of Wisteria and other millettioid-phaseoloid genera using data from phytochemical and embryological analyses as well as noting the chromosome count of 2n =16 in Wisteria compared to those of other genera whose members frequently have 2n = 22 or 2n = 24. It is notable that within the Callerya group, two species of Afgekia; A. sericea and A. mahidoliae also have chromosome counts of 2n = 16 (Prathepha 1994;Prathepha and Baimai 2003). Zhu's (1994) concept of Tribe Wisterieae thus was based solely on four samples of Wisteria and one accession of Millettia [Callerya] reticulata (Table 1).
One taxon recently recognised as belonging in the Callerya group was included under the names Wisteria japonica or Millettia japonica (Doyle et al. 1997;Doyle et al. 2000;Kajita et al. 2001;Hu and Chang 2003). The inclusion of this taxon in Wisteria was originally based on the deciduous leaves, twining habit, pendulous inflorescences and flowers where the wing petals are free from the keel (Siebold and Zuccarini 1839;Bailey and Bailey 1949;Geesink 1984;Iwatsuki et al. 2001;Compton and Thijsse 2013;Compton 2015c). Its summer flowering habit, paniculate inflorescences and absence of callosities on the standard petals have also been used to segregate it from Wisteria (Gray 1858;Dunn 1912a;Valder 1995).
All genera currently comprising the Callerya group (Table 1): Afgekia, Callerya, Endosamara, Wisteria [incl. Millettia japonica)] and Sarcodum possess bracts enclosing the apical floral buds prior to anthesis and all bear either true racemes or true panicles. All are lianas with the exception of two tree species Callerya atropurpurea (Wall.) Schot and C. vasta (Kosterm.) Schot. The genus Sarcodum was not included in the analysis of Jansen et al. (2008) and has not been sampled for DNA analysis prior to this paper but the generic morphological characters (Table 4) place it firmly within the Callerya group.
The genus Callerya Endl., the largest genus within the group with 33 species (Table  1) has subsequently been found to be polyphyletic (Li et al. 2014). Without a comprehensive morphological study of the genus and its near relations and in the absence of additional DNA evidence, it is fair to state that the genus has been something of a catch-all for taxa that bear some morphological affinities with each other Lôc 1996). The purpose of this paper therefore is to test generic boundaries within the Callerya group by reassessing morphological characters and by a comprehensive molecular sequencing investigation of representative species of all taxa within the group using both nuclear and chloroplast genes.

Molecular preparation and sequencing
Taxon sampling included those taxa in the DNA based phylogenies of Doyle et al. (1997), Hu et al. (2002), Hu and Chang (2003), Jansen et al. (2008) and Li et al. (2014 -but not including several species they recognised in the Callerya cinerea complex). Three chloroplast regions were included in the study. Two protein coding genes: matK and rbcL, and the intergenic spacer ndhJ-trnF. One nuclear gene region was also included in the study ITS1, 5.8S and ITS2. Fresh DNA was extracted from the previously unsampled Sarcodum scandens (Tables 1,2). For the ITS dataset, 12 additional sequences representing the Callerya group and 26 outgroup sequences were included from GenBank; the matK dataset comprised an additional five Callerya group sequences and 17 outgroup sequences from GenBank and the rbcL dataset, a further two ingroup and 12 outgroup sequences from GenBank (Table 2). Millettia japonica has also been confused with Wisteria floribunda in DNA sampling (see GenBank KT119544) and as a result of this, we have chosen to include three different verified samples in this study in order to ascertain its placement within the Callerya group.
Outgroup taxa (Table 2) for each analysis comprised several accessions that represented taxa from other Tribes within the IRLC e.g. Hedysareae, Galegeae and Trifolieae (Lewis et al. 2005) and several from outside the IRLC, e.g. Robineae, Loteae, Sesbaneae, Millettieae, Abreae, Phaseoleae and Indigofereae. The genus Schfflerodendron Harms was selected as the outgroup upon which to root all trees owing to its position in the LPWG (2017) RAxML Maximum Likelihood analysis. Its position as sister to Callerya atropurpurea -and these two to Glycyrrhiza -in turn all sister to the Callerya group and the rest of the IRLC suggest that it is the most appropriate candidate for choice as outgroup. Additional outgroup sequences were generated of W095 (see codes to samples, Table 2) Schefflerodendron usambarense (Tribe Millettieae), W113 Lotus uliginosus (Tribe Loteae) and W115 Austrosteenisia glabristyla (Tribe Millettieae) with the addition of 14 other legume genera from GenBank representing additional Tribes all of which sit outside the IRLC (Table 2).
We generated 49 sequences of the nrDNA ITS spacer region, including one for the outgroup taxon Schefflerodendron usambarense. Sequence data was also generated for three plastid markers: 51 Callerya group sequences from the ndhJ-trnF cpDNA intergenic spacer, 53 sequences from the matK gene and 57 sequences from the rbcL gene. Sequences of three outgroup taxa (i.e. Austrosteenisia glabristyla, Lotus uliginosus and Schefflerodendron usambarense) were also obtained for these three plastid markers (see Table 2). Summary statistics of support levels at critical nodes of all trees generated in this study (Suppl. material 1 Figs S1-S6), derived from Maximum Likelihood (ML) analysis and Bayesian inference (BI) analysis, are shown in Table 3.
The DNA extraction protocol for all 54 samples (with numbers from W002 to W115) and the seven samples labelled W1, W2, W3, W5, W6, W8 and W10 (Table 2) used a modified CTAB protocol (Doyle and Doyle 1987). DNA extraction from herbarium specimens followed the protocol used by Särkinen et al. (2012) with some minor amendments.
For all accessions ( Table 2) the gene rbcL was amplified with primers 1F (Fay et al. 1997) and 1460R (Fay et al. 1998). To amplify degraded and/or low quality DNA two overlapping shorter fragments were amplified with the original primers and internal primers 636F and 724R (Fay et al. 1997). PCR reactions for all primer combinations were performed in 25μl volumes containing final concentrations of 1× Bioline Biomix Red, 0.35μM of each primer, 0.2mg/ml BSA (bovine serum albumin), and 20ng DNA template. Cycling conditions for the reactions using primers 1F and 1460R were 94 °C for 120s, then 30 cycles of 94 °C for 60s, 51 °C for 30 s, 72 °C for 120s, and finally 72 °C for 7 minutes. For the shorter fragments this protocol was modified by decreasing the elongation time to 90s and increasing the number of cycles to 40 for weaker amplicons.
Again for all accessions, the intergenic spacer ndhJ-trnF was amplified with the primers ndhJ and TabE using the PCR protocol listed in Shaw et al. (2007). Low quality or degraded DNA necessitated the utilisation of primers that amplified two shorter, overlapping segments of this region. We designed two additional primers internal to the ndhJ-trnF spacer in order to overcome this problem: 456F -ATGGGC-CGGATTCTATTTGT and 725R -TGATTAGTGGTCTAGATCATCAT. The PCR protocol for the shorter fragments was the same as above, apart from increasing the number of cycles to 40 for weaker amplicons.
For all accessions the nrDNA Internal Transcribed Spacers (ITS1 and ITS2) were amplified with primers ITS4 and ITS5 (White et al. 1990;Baldwin et al. 1995) or with 17SE and 26SE (Sun et al. 1994 Sequencing of 44 taxa for ITS and 54 taxa for ndhJ-trnF, matK and rbcL were performed at GATC Biotech (www.gatc-biotech.com; Konstanz, Germany). Table 2. Vouchers of taxa used in the phylogenetic analyses. Included are all the Callerya group taxa as well as all outgroup taxa used (marked x in the last column).
The W numbers represent taxa sampled in this analysis for one or more of three plastid genes (rbcL, ndhJ-TabE and matK) and/or the ITS nuclear spacer regions and G numbers are additional sequences of both ingroup and outgroup taxa downloaded from GenBank. For the seven accessions labelled W1 to W10 (see Table 2) DNA extractions used a similar protocol to that mentioned above. PCR amplifications were performed using the same primers as already mentioned for nrDNA ITS, cpDNA matK and cpDNA rbcL with the following different protocol: PCRs were performed in 25 μL volumes, containing 12.5 μL DreamTaq PCR Master Mix (2×) (4 mM MgCl2; Thermo Fisher Scientific, Waltham, MA, USA), 0.5 μL of each primer (100 ng μL−1), and 1 μL DNA template. TBT-PAR [trehalose, bovine serum albumin (BSA) and polysorbate-20 (Tween-20)] was added to reduce the inhibitory effects of polysaccharide and phenolic compounds (Samarakoon et al. 2013).

ndhJ-TabE
For the accessions W1 to W10 (Table 2) all amplifications were performed on a 9700 GeneAmp thermocycler (ABI, Warrington, UK). All PCR products were purified with either the QIAquick PCR kit (Qiagen, Hilden, Germany) or the Nucleospin Extract II kit (Machery-Nagel, Düren, Germany), following the manufacturers' protocols. Cycle sequencing reactions were performed in 5 μL reactions using 0.5 μL Big-Dye Terminator cycle sequencing chemistry (v3.1; ABI) and the same primers as for PCR. Complementary strands were sequenced on an ABI3730 automated sequencer.

Phylogenetic procedures and analyses
Sequences of each region were edited and compiled in Geneious (version 8.1.9; Kearse et al. 2012) and aligned with the MUSCLE algorithm (Edgar 2004) implemented in AliView (Larsson 2014). The ends of the alignments were trimmed to the point where all sequences were present and base calls were unambiguous.
Phylogenetic analyses were conducted on the plastid, ITS and combined plastid/ ITS matrices using two approaches, Maximum likelihood (ML) and Bayesian inference (BI). For the ML approach, we used the software RAxML (v. 8.2.8;Stamatakis 2014) as implemented on the CIPRES portal (www.phylo.org) with 1,000 rapid bootstrap replicates followed by the search of the best ML tree; the GTRCAT model was used and all the other parameters were set as default settings. The Bayesian Markov Chain Monte Carlo (MCMC) approach was performed using the software MrBayes (version 3.2.6; Ronquist and Huelsenbeck 2003) as implemented on the CIPRES portal. The best-fit DNA substitution model was tested using JModelTest 2 (version 2.1.6; Darriba et al. 2012) as implemented on the CIPRES portal. The General Time Reversible (GTR) model with a proportion of invariable sites and a gamma shape to account for rate heterogeneity among sites (GTR+I+G) was selected for both partitions. The analyses were run twice each for 10 million generations and sampled every 1000 th generation. The MCMC sampling was verified using Tracer (Rambaut and Drummond 2009) and was considered adequate when the effective sampling size was higher than 200. A burn-in period of one million generations was applied to each run. The remaining trees from both runs were compiled using the ''allcompat'' option in MrBayes to produce a maximum credibility tree with Bayesian posterior probabilities (BPP) for each node. In both combined ML and BI analyses, the plastid and ITS partitions were allowed to have partition-specific model parameters. Schefflerodendron usambarense was designated as outgroup taxon in all analyses. Support values for nodes of critical taxa in the Discussion are shown in Table 3.

Morphological study
The morphological key to the species was based on examination of living material in cultivation in UK and USA and in the wild in China, Japan, Laos, Myanmar, Thailand and Vietnam. Herbarium specimens were examined including the collection of all relevant genera in the Callerya group at K and BM. Online collections were examined at the Chinese Virtual Herbarium, CVH (http://www.cvh.ac.cn/en); JSTOR Global Plants (https://plants.jstor.org/); Herbarium, Muséum National d'Histoire Naturelle, Paris, MNHN (https://science.mnhn.fr/institution/mnhn/collection/p/item/search/ form?lang=en_US); Herbarium Royal Botanic Garden, Edinburgh, RBGE (http://data. rbge.org.uk/search/herbarium/) and Nederlandse Natuurhistorische Collecties, Naturalis (http://bioportal.naturalis.nl/). See Appendix 1 for a full list of all specimens used as the basis for the new generic descriptions. Herbarium acronyms follow Thiers (2019, http:// sweetgum.nybg.org/science/ih/). A full list of all Herbaria cited is found in the acknowledgements. A list of the critical characters measured for this study is shown in Tables 4, 5.

Results
New diagnoses (emended where necessary) -and full descriptions -are given for all genera in the taxonomic treatment section, because nearly all established genera have been modified over various historical treatments to include and/or exclude species such that their present concepts are often significantly different from the original protologue. Keys to genera and to all species (excluding those Chinese taxa of Callerya s.str. that we were unable to access) and extensive synonymy and typifications are also provided.
The combined analyses are consistent with respect to their ingroup topologies and are combined in the reference phylogenetic tree of this paper, Fig. 1 (see also Suppl. material 1: Figs S1, S2). In the plastid analyses (Suppl. material 1: Figs S3, S4) the main difference to Fig. 1 is Serawaia grouping with Clades D + E with no support in the BI analysis -and with Clades C + D + E, with no support, in the ML analysis. The ITS BI analysis is similar to Fig. 1 except for the merging of Clades B + C. In the ITS ML analysis Clade A also breaks down together with Clades B + C as Endosamara is attracted into Clade C from Clade A. As an indication of variability across the four genes, the average percentage identity over the alignments (i.e., pairwise percentage of identity) is: plastid vs. ITS analyses (92.5%, 77.1%) and within the plastid analyses, matK, rbcL and ndhJ-trnF (95.4%, 96.9% and 86.6%). Table 3. Summary of support values at critical nodes for trees derived from the six phylogenetic analyses (Suppl. material 1: Figs S1-S6). Levels of ML Bootstrap (BS) -and Bayesian Inference (BPP) -support are listed for each of the combined plastid and nuclear, plastid only and ITS analyses, for genera and clades discussed in the text. Rows in bold represent genera. Single accessions are where only one sequence was available for a taxon and comments are included in the table highlighting those conflicting arrangements of taxa between the plastid and ITS analyses. BS and BPP support of 85%/0.95 and higher are considered strong, 65-85%/0.9-0.95 as moderate, and below 65%/0.9 as weak.

Outgroups
The Callerya group sensu ; Lôc and Vidal (2001);  and Sirichamorn et al. (2016) is not supported in its entirety in our analyses but rather, what emerges are four elements comprising the IRLC, each fully supported here ( Fig. 1; Table 3), i.e. Tribe Wisterieae, Glycyrrhiza, Adinobotrys and a clade containing the Galegeae s.l., Hedysareae, Cicereae, Fabeae and Trifolieae s.l. (henceforth the "Temperate Tribe block"). Glycyrrhiza (Bootstrap or BS 100%; Bayesian Posterior Probability or BPP 1) is not supported as sister to Tribe Wisterieae and neither is Adinobotrys (BS 100%; BPP 1), which, in addition, is not supported to have a position within Tribe Wisterieae either. Adinobotrys is thus reinstated as a genus here and removed from the Callerya group. The positions of Tribe Wisterieae, Glycyrrhiza and Adinobotrys remain equivocal as regards their sister group relationships to the rest of the Temperate Tribe block of the IRLC.
The Callerya group i.e. Tribe Wisterieae without the genus Adinobotrys comprises five strongly supported clades with the first two in a basal grade leading to Clades C + D + E. The crown node of the tribe is fully supported in both the combined and plastid analyses (BS 100%; BPP 1) and in the ITS BI analysis (BPP 1), although only weakly so in the ITS ML analysis (Fig. 1, Table 3, BS 61%). Clade A (fully supported in the combined and plastid analyses but weakly so in the ITS analyses [BS 59%, BPP 0.86]) contains the genus Sarcodum (BS 100%; BPP 1), sequenced and analysed for the first time here, which is sister to Endosamara (BS 100%; BPP 1) and the new monospecific genus Sigmoidala (BS 99%; BPP 0.99) described in this paper. Clade B comprises two new genera described here, Nanhaia (BS 100%; BPP 1) and Wisteriopsis (BS 91%; BPP 1) and both are fully supported as Clade B ( Fig. 1; Table 3). Our results confirm that the incorrectly attributed Millettia japonica is strongly supported within our new genus Wisteriopsis. Wisterieae. The tree is derived from the combined plastid and ITS, RAxML bipartitions analysis representing 77 (36) ingroup samples (taxa) and 59 (40) outgroup samples (taxa). The outgroup Schefflerodendron is used to root the trees. Lines in bold on the phylogeny incorporate results from the combined Bayesian Inference analysis, demarcating clades with BPP (0.95) support and above. Nodes are marked up with bootstrap values as percentages derived from the combined ML analysis with values of 50% or less marked in red. The collapsed portion of the tree, below the IRLC and above Schefflerodendron, represents the following genera (see Suppl. material 1: Figs (Wisteria). Each clade is further subdivided to represent the genera (except for the single accession of Serawaia which is incorporated with Whitfordiodendron in Clade C2) and E1 and E2 represent the geographical disjunction of species in Wisteria. Outgroups within the IRLC in purple include Glycyrrhiza, Adinobotrys and representatives of the Temperate Tribe block. The ingroup (IRLC) and Tribe Wisterieae are demarcated with arrows on the tree. Clade C is strongly supported (BS 87%; BPP 0.98) in the combined analyses but is more labile with some genera excluded and others included in the plastid and ITS analyses ( Fig. 1; Table 3). A much reduced Callerya s.str. together with C. bonatiana is strongly supported in the combined and plastid BI analyses (BPP 0.95; 0.96), but only weakly so in the ML analyses. The single accession of C. bonatiana has no support for grouping with Callerya in the ITS analyses. Callerya above C. bonatiana is strongly supported in the combined ML and BI analyses (BS 100%; BPP 0.95). The grouping of Afgekia (BS 100%; BPP 1), the resurrected genus Whitfordiodendron (BS 100%; BPP 1) and the two new genera described here, Kanburia (BS 100%; BPP 1) and Serawaia (single accession), is moderately supported in the combined ML (BS 72%) and well supported in the combined BI (BPP 0.99) analyses. This grouping breaks down in the plastid analyses and is weakly supported in the ITS analyses (Table 3). Serawaia is strongly supported within Clade C in the combined analyses (BS 87%; BPP 0.98) in a position (Fig. 1, Table 3), with no support, as sister to Whitfordiodendron. It is in an unresolved position in the plastid analyses and is weakly supported as sister to Kanburia in the ITS analyses (Table 3).
Clade D comprises two genera, Padbruggea (BS 91%; BPP 1) which is reinstated as a genus here and Austrocallerya (BS 98%; BPP 1), a new genus described here. Our results reveal that Afgekia filipes belongs in our reinstated genus Padbruggea and the transfer back is made in this paper. The two genera are also strongly supported together as Clade D (BS 98%; BPP 1). Clades D + E are strongly supported in all analyses (combined ML [98%] & BPP [1]; plastid BPP [0.97] and ITS ML [90%] & BPP [1]), but in the plastid ML analysis support is weak (BS 68%). Finally Clade E comprises Wisteria (BS 100%; BPP 1), with two North American taxa fully supported as sister to the three Asian species of the genus (BS 100%; BPP 1). The relationship of W. brachybotrys as sister to W. floribunda and W. sinensis is also fully supported (BS 100%; BPP 1).

Morphology of the Callerya Group
Schot (1994) segregated her species of Callerya into two groups based on the presence or absence of stipels and, when present, whether they were persistent or caducous. We have found no evidence that stipel presence or absence has any taxonomic significance in Calleya s.l. Lôc (1996) and  segregated species on the basis of the presence or absence of an indumentum on the dorsal surface of standard petals. We concur with Lôc (1996) and  but in addition regard the inflorescence type and various floral, fruit and seed types to be equally significant in delimiting taxa (see the Key to the Genera and Table 4 for a list of significant characters). The key to fruiting specimens of Callerya s.l. (Lôc 1996: 56) emphasised stipellae characters as distinctive, an observation for which we find no support.
Using our revised generic concepts and species assigned to them ( Table 1) and comparison of morphological characters (Table 4), the synapomorphies diagnosing the Callerya group are: the lianescent habit (except for the tree species Adinobotrys atropurpureus and A. vastus); flowers inserted singly on the axis in either axillary or terminal racemes and/or panicles, and bracts either fully or largely enclosing the flower buds at the inflorescence apex, which are usually longer and often wider than the buds. There are, however, some exceptions. Floral bracts are caducous at anthesis in most of the Callerya group except in Adinobotrys atropurpureus, Nanhaia speciosa, Serawaia strobilifera and Wisteriopsis where they are persistent.
Gibbosities, which are small protuberances that develop beneath the leaf pulvinus above the stipule where it is attached to the stem, are absent in most of the Callerya group but are present in both Wisteriopsis, Nanhaia and Serawaia (Table 4; Fig. 2R-S). Bracteoles may be found either at the base of the calyx or along the pedicel. They are present in most genera but absent in Endosamara, Sigmoidala, Kanburia and Afgekia. They are also absent in Wisteria frutescens (Table 4).
Genera in the Callerya group often differ from each other (Table 4, Fig. 2) according to the presence of callosities at the base of the standard petals. Callosities occur in five distinct types: a) Boss callosities form two slightly raised domes or swellings on either side of the midline of the standard lamina, at the point of its upward flexion above the claw ( Fig. 2A). The standard in the latter case often appears to be smooth but the bosses hold the two wing petals close to the standard prior to anthesis. Boss callosities are found in Adinobotrys, Endosamara, Sigmoidala, Sarcodum, Nanhaia, Serawaia and Wisteriopsis and occasionally in Callerya and Whitfordiodendron; b) Arched callosities are paired half-moon or crescent shaped arches forming ridges of hardened tissue that curve up from the base towards the midline over the staminal sheath (  There are notable differences in the fruits and seeds among the genera. In Endosamara and Sarcodum the exocarp separates from the endocarp and some degree of separation also occurs in Wisteriopsis. In Endosamara the pods are clearly septate with transverse walls between each seed, forming loments (see Endosamara racemosa in Geesink 1984: 63 Pl. 1, 5;Lôc and Vidal 2001: 17, Pl. 3). In Sarcodum the sausageshaped or botuliform pods which initially have a fleshy exocarp, are also fully septate but do not form loments (see Sarcodum scandens in Geesink 1984: 63 Pl. 1, 7;Lôc and Vidal 2001: 9, Pl. 1). The North American Wisteria frutescens has nonseptate pods. In all other genera in the Callerya group the endocarp is subseptate with seeds making indentations in the surrounding pith while areas between the seeds appear as irregular, often indistinct transverse septa. In Afgekia the funicle as well as the hilum (see Endlicher (1843) first described the genus Callerya based on Marquartia tomentosa Vogel, which had been described that same year from southern China (Vogel 1843: 37   Vogel's illustration clearly shows the leaves with persistent stipules, each with five leaflets, long persistent floral bracts and densely sericeous ovary, all characters diagnostic of that species. This meant that not only the generic name Marquartia but also Callerya tomentosa (Vogel) Endl. had to be replaced according to Art. 11.4 of the ICN (Turland et al. 2018). The replacement Callerya nitida (Benth.) Geesink was published formally by Geesink within his revision of Tribe Millettieae (Geesink 1984: 82). Callerya nitida is thus the type species of the genus Callerya. Endlicher's description of Callerya mentioned the compressed, woody and leathery pod which he stated contained either a few seeds or a single seed which was ovoid-circular and flattened-compressed. Callerya nitida ( Fig. 2L-N) does indeed possess flattened pods with up to ten compressed seeds (see Schot 1994: 28, Fig. 3). The species may also become a scandent shrub but is not arborescent as Endlicher implied. Geesink (1984) recognised that species of Callerya had true paniculate inflorescences. He also sank three genera; Adinobotrys Dunn, Padbruggea Miq. and Whitfordiodendron Elmer into his redefined Callerya and transferred two Sections created by Dunn (1912a) from Millettia -Sect. Eurybotryae Dunn and Sect. Austromillettia Dunn -into Callerya and created a new monotypic genus Endosamara Geesink from Sect. Bracteatae Dunn based on Robinia racemosa Roxb. (1832).
Callerya was revised by  and was treated as belonging in Tribe Millettieae (see Table 1). Schot recognised 19 species from China, south-east Asia and Australasia and since then many more species have been described, bringing the total number to 33 (Sirichamorn et al. 2016). All species sensu Schot (1994) except C. atropurpurea and C. vasta, are vigorous climbing or scandent woody shrubs. Inflorescences are paniculate with either axillary racemes or secondary panicles and frequently possess rather thick to woody inflorescence axes with prominent bud scars. Floral bracts are generally short and can be narrow or broad according to species and are in most species caducous, rarely persistent. Flowers may be white, green, red, brownish-yellow, lilac, pink or deep purple. In the type species Callerya nitida, the wings are distinctly shorter than the keel. Fruits, which can be either flattened or inflated are velutinous and ribbed, occasionally wrinkled or smooth and the seed chambers are subseptate. Seeds 2-9, large, ovoid to ellipsoid (Tables 1, 4).  2) chronicled the transfer of species from other genera into Callerya and included eleven species from Millettia, the first two of which were Pterocarpus australis Endl. Prod. Fl. Norfolk (Endlicher 1833: 49) and Pongamia atropurpurea Wall. (Wallich 1830: 70).
Although the genus Pterocarpus is placed in Tribe Dalbergieae (Klitgaard and Lavin 2005), the species P. australis has been shown to belong in the Callerya group (Li et al. 2014). Pongamia Adans. is now treated as being synonymous with Millettia, with the type P. pinnata (L.) Pierre being transferred to Millettia pinnata (L.) Panigrahi by Panigrahi and Murti (1989). Pongamia atropurpurea which was transferred into Callerya by Schot (1994: 15) has also been found to belong in the Callerya group (Li et al. 2014).  further transferred the Australian species originally described as Wisteria megasperma F.Muell. (1858) into Callerya. This too has been confirmed to belong within the Callerya group (Li et al. 2014).
In their analyses using combined data from chloroplast trnK and matK sequences, Hu et al. (2000) showed that Callerya reticulata was sister to a clade supported by BS 100% with Wisteria frutescens and W. sinensis. In a later paper using sequence data from nuclear DNA ITS spacers and a larger sampling of Callerya and Wisteria as well as Afgekia filipes, Hu et al. (2002) found that Callerya was polyphyletic occurring in four different clades. Wisteria frutescens was strongly supported sister to W. brachybotrys, W. sinensis and W. floribunda with BS 100%. The Wisteria clade was sister to a clade with Callerya megasperma, C. australis and Afgekia filipes with strong BS (85%) support. In a later analysis Hu and Chang (2003) using the more conserved rbcL chloroplast gene, found that Callerya vasta was early branching to a clade of Wisteria sinensis sister to Afgekia sericea while Endosamara racemosa was sister to Millettia japonica.
Schot included nine synonyms within her concept of C. cinerea, a species that she recognised to have a wide distribution from Nepal in the west to the Chinese coast in the east (Schot 1994: 17). In our analyses we have utilised two specimens of Callerya cinerea, one from Thailand, the other from China, ( Table 2) but these sheets lack diagnostically significant fruiting material and may not equate fully with the holotype material seen at Kew from Bangladesh, Sylhet, Chittagong, (Wallich 5888; K000881022). Within the C. cinerea complex, leaflet number, pod thickness and seed shape and number appear to be important characters.  split C. cinerea into groups of species based on leaflet number: 3-5 in C. tsui, C. dorwardii and C. sphaerosperma and 5-7 in the other seven species.  also segregated the species on the degree of pod inflation: flattened with lenticular seeds in C. congestiflora, C. dielsiana and C. longipedunculata; inflated with globose seeds in C. cinerea, C. dorwardii, C. gentiliana, C. oosperma and C. sericosema. We have only been able to sample material of two of the resurrected species recognised by  that were included in C. cinerea by , namely C. dielsiana and C. oosperma. A further investigation of this group of Chinese species is needed to fully assess species delimitations. If all these species belong together with C. nitida and C. cochinchinensis as indicated both by Li et al. (2014) and from our preliminary results here, it seems that Callerya s.str. might comprise as many as twelve species. Reference to the unwinged nature of the fruit was in comparison to some species of Pterocarpus Jacq. (Tribe Dalbergieae) whose fruits have a distinct wing-like exocarp. Miquel also noted that he had not seen mature fruits.
Our examination of the species described as P. dasyphylla Miq. (1855), revealed that the pods were readily distinguished from others in the Callerya group by their inflated but broadly flattened-cuboid shape with distinct longitudinal ridges and furrows and by the 1 or 2 compressed obovoid seeds possessing long strap-shaped hila 18-36 × 4-7 mm (Table 5).
In his protologue of Adinobotrys filipes, Dunn (1911: 196) noted that in its appearance the species had more slender pedicels than those in what he considered to be the closely related A. erianthus (here in Whitfordiodendron) and that it approached Padbruggea in its auriculate standard (Dunn 1911). Dunn's latter reference may also refer to the papillate callosities present on the standard of A. filipes but which are absent on the smooth standard of A. erianthus. In her monograph on Callerya, Schot (1994: 3) commented:

Craib (1928) argued that Padbruggea and Whitfordiodendron were congeneric. He based his arguments on the intermediate position of Adinobotrys filipes Dunn (now Afgekia filipes Geesink). This species resembles in habit mostly Padbruggea dasyphylla, but has the generic characters of Adinobotrys.
Craib recombined Adinobotrys filipes in Padbruggea, along with a good measure of uncertainty as to whether he believed the species really belonged in that genus or in Adinobotrys (Craib 1928: 397). He also postulated that Elmer's Whitfordiodendron may belong in Padbruggea thereby highlighting the morphological difficulties with respect to these taxa faced by later workers such as Geesink (1984).
Schot in her synonymy of Callerya dasyphylla also included Milletia oocarpa Prain, distinguished from P. dasyphylla by its ovoid as opposed to compressed obovoid fruits and M. maingayi Baker which differs in its more numerous, smaller and more densely tomentose leaflets (Schot 1994: 20). We have recognised this as Padbruggea maingayi in this paper (see below).
The status of Afgekia filipes has long been debated as it has true panicles as opposed to racemes and shorter calyx teeth than those of the other two species of Afgekia (see Table 5). It also has a single pair of papillate callosities on the standard as opposed to two pairs found in both A. sericea and A. mahidoliae. Afgekia filipes has entirely glabrous anthers as opposed to anthers with a basal tuft of hairs and it has much larger fruits and seeds (Table 5). It was originally described as Adinobotrys filipes Dunn on the basis of its large single seeded pods (Dunn 1911: 195). Geesink (1984: 76) transferred Adinobotrys filipes into Afgekia adding: the general habit, the shape of the calyx, and the glabrous anthers are indeed similar to certain species of Padbruggea. It differs in the absence of bracteoles and the long pedicels. The pods were unknown until 1975, but then it appeared that the seeds showed an elongated fleshy funicle with a corresponding elongated hilum. Geesink (1984: 76) concluded that the morphology of A. filipes indicated that it was a less derived species than A. sericea and A. mahidoliae and alluded to its affinities with Padbruggea within which it had been placed by Craib (1928) along with three other species in Adinobotrys.
In his transferral of the species into Afgekia, Geesink noted the apparent absence of bracteoles, the length of the pedicels and the elongated hilum on the seeds (Geesink 1984: 77). Both Lôc and Vidal (2001) and  followed Geesink, maintaining the species in Afgekia. Sirichamorn (2006) examined 37 living specimens of A. sericea, 50 specimens of A. mahidoliae and 32 specimens of A. filipes from wild material in Thailand and found that A. filipes posseses bracteoles, that pedicel length among the species overlaps and that the overall size of the seeds are three times that of the other two species of Afgekia, i.e. c. 80 mm vs. 15-25 mm long (Tables 4, 5). Prathepha and Baimai (2003) mentioned the existence of Afgekia filipes in their RAPD and nucleotide sequence analyses of A. mahidoliae and A. sericea, although they did not state their reason for excluding the species. Sirichamorn (2006), based on morphometric and molecular data, clearly showed that Adinobotrys filipes does not belong with Afgekia (Table 5).
We have examined material of both Afgekia filipes and Callerya dasyphylla and agree that there are indeed similarities between the two species. We have confirmed Sirichamorn's (2006) discovery that Afgekia filipes does have short, linear bracteoles that are attached at the base of the calyx (Table 5). Both species possess inflated fruits with a velvety indumentum and oblique longitudinal ridges and furrows but those of C. dasyphylla are broader and flatter, with the dorsal midline flanked by two large folds or flanges that meet at the apex. Our results show that Afgekia filipes, originally described by Dunn (1911) in Adinobotrys, belongs in the genus Padbruggea and it is reinstated in that genus here (see Taxonomic treatment below) following Craib (1928). The diagnostic characters of Padbruggea filipes are shown in Table 5. Whitfordiodendron Elmer. Leafl. Philipp. Bot. 2: 689, 743 (1910) Elmer (1910) in his protologue of the illegitimate but valid name Whitfordia scandens stated that he had only seen young not mature fruits but that they were "thick, hard, canescently velvety and 1-seeded". He also noted the puberulent dorsal surface of the standard petal of the deep red flowers (Elmer 1910: 691). The generic name Whitfordia was already utilised for the fungal genus Whitfordia Murrill (Murrill 1908: 407), a synonym of Amauroderma Murrill (Murrill 1905: 366) and although Whitfordia Elmer (with W. scandens) was described entirely in English, together with that of the transfer to Whitfordiodendron as an erratum in an appendix to the same volume, the name is nevertheless still valid. Under the International Rules of Nomenclature adopted in Vienna in 1905 and Brussels in 1910, a Latin diagnosis was a requirement for valid publication of a name of a new taxon on or after 1 January 1908 (Art. 36). As a result of discrepancies and disagreements between the American Code of Botanical Nomen-  Merrill [1934: 159] who independently concluded that Art. 38 of the Cambridge Rules validated Elmer's Whitfordiodendron), Merrill proceeded to describe the new species Whitfordiodendron sumatranum Merr., which he stated was close to W. myrianthus (i.e. to W. nieuwenhuisii (J.J.Sm.) Dunn). Merrill (1934: 160) also made what he believed to have been four new combinations in Whitfordiodendron but he was evidently unaware that W. atropurpureum, W. erianthum, W. myrianthum and W. nieuwenhuisii had already been combined in Whitfordiodendron by Dunn (1912b: 364).
Our morphological examination has revealed that two species previously included within Callerya share a suite of characters with Whitfordiodendron scandens. The most notable characters are: a) the flowers borne on extremely short pedicels 0.5-3 mm long vs. (2-)3-8 mm long in Callerya s.str.; b) the inflated, ovoid, rugose to ridged or ruminate pods with 1-3 seeds (if more than one-seeded then these often becoming fused together, Fig. 2P-Q) vs. pods flattened or if inflated then convex around seeds and contracted between them, the seeds being separate in the pod in Callerya s.str.; and c) most significantly, the sericeous keel petals which are particularly densely hairy along their lower margins (keel glabrous in Callerya s.str., see Table 4).
Based on nrDNA ITS sequence data, Li et al. (2014) showed that Callerya eriantha, C. scandens and C. nieuwenhuisii formed a clade with BS (100%) which is sister to C. eurybotrya and C. reticulata.

"affinis Millettieae Wight et Arn. sed ovario stipitato, legumine monospermo indehiscente differt [related to Millettiae Wight & Arn. but differs by having a stipitate ovary and indehiscent one-seeded pod]".
Dunn made a further distinction between Padbruggea (which he understood to comprise P. dasyphylla and P. maingayi) and Adinobotrys, stating that the inflorescence in Padbruggea was lax and that Padbruggea lacked any appendages on the wings and keel petals (Dunn 1911: 197). Dunn (1911) included five species in Adinobotrys without assigning any one of them as the type species; A. erianthus (Benth.) Dunn, A. filipes Dunn (see above), A. nieuwenhuisii (J.J.Sm.) Dunn, A. myrianthus Dunn and A. atropurpureus (Wall.) Dunn. Geesink (1984: 83) typified Adinobotrys on the species A. atropurpureus, the only species of the five which is a tree and not a liana. The following year Dunn (1912b) added A. scandens (Elmer) Dunn in the belief that Elmer (1910) had not validly published the name under Whitfordiodendron. Dunn, recognising the uncertainty of the validity of Whitfordiodendron, validated Elmer's names firstly in Whitfordiodendron (including W. scandens) and then into his new genus Adinobotrys (Dunn 1912b: 364, 365).  included A. atropurpureus, A. erianthus and A. nieuwenhuisii in Callerya and recognized A. myrianthus as conspecific with C. nieuwenhuisii. Schot did not include A. filipes in Callerya as she treated the species as belonging in Afgekia (Schot 1994: 3).
Adinobotrys has several morphological characters that separate it from the other allied genera within the Callerya group; trees vs. lianas, stipules 2-4 mm long, floral bracts short, c.1-3 mm long, standard petal glabrous (although this is not unique to Adinobotrys), pods inflated with glabrous, rugose surfaces and large ovoid seeds with short elliptic or circular hila (see Geesink 1984: 64 Pl. 2, 14).
Results from sequence data of nuclear ITS and chloroplast matK showed that Callerya atropurpurea was placed sister to the rest of the Callerya group (Li et al. 2014).
Callerya group taxonomy 2: Additional genera within the Callerya group as defined in treatments prior to this study.

Sarcodum Lour., Fl. Cochinch. 2: 462 (1790)
The Portuguese Jesuit missionary and botanist João de Loureiro was the first to describe the genus Sarcodum in 1790 based on its seed pods which are fleshy when young [sarcos = Gk fleshy] (Plate 2G). The type species S. scandens was described as having rose-coloured flowers in simple spikes and was found growing in woods of Cochinchina, i.e. modern day Vietnam. It has since been collected in China on Hainan Island, in Indonesia and in the Philippines. In 2017 it was discovered by co-author S. Mattapha in Bolikhamxai Province in Laos. Sarcodum scandens has standard petals 10-13 × 6-8 mm and leaves with 17-45 leaflets. Two further species have recently been described; S. bicolor in 1999 with standard petals 13 × 8 mm and leaves with 9-15 leaflets from Sumba in the Lesser Sunda islands of Indonesia and S. solomonensis in 2008 from the Solomon Islands with standard petals 6 × 5 mm and leaves with 17-27 leaflets (Clark 2008: 156, Table 4).
Distinguishing characters are the many small sericeous, elliptic leaflets; long stipules; long caudate floral bracts; racemose inflorescences; flowers with campanulate calyces and five very short, acute teeth; glabrous standard petals with boss callosities, elongating persistent styles on the developing pods post anthesis, and fleshy, cylindrical, botuliform fruits becoming hard when mature. The glabrous exocarp dries to dehisce from the septate tan-coloured chartaceous endocarp in which lie the 4-10 ellipsoid to reniform seeds (Kirkbride et al. 2003, Table 4).
None of the three species of Sarcodum have been included in any DNA based phylogeny prior to this study, although its affinities with other genera in the Callerya group have been postulated (Geesink 1984;Lavin et al. 1998).

Endosamara R.Geesink Leiden Bot. Ser. 8: 93 (1984)
The unique monospecific genus Endosamara has never been included in Callerya but is nevertheless considered to be a close relative (Geesink 1984: 94). The Scottish surgeon and botanist William Roxburgh spent several decades in India where he described Robinia racemosa, a climbing shrub from the Circar Mountains [Eastern Ghats] north of Madras, which had rose-coloured flowers in what he described as racemes but we now know to be panicles (Roxburgh 1832: 329). The species was introduced to the Calcutta Botanic Garden by Henry Colebrooke in 1803 from the Coromandel region of south-east India (Roxburgh 1814: 56). The single species has a widespread distribution across India, Laos, Myanmar, Philippines, Sri Lanka and Thailand. It was placed in Millettia by Bentham (1853) and was later recombined in Wisteria (Dalzell in Dalzell and Gibson 1861: 61). Dunn (1912a: 135) placed Millettia racemosa in his monotypic Sect. Bracteatae, a taxon which was later recognised as Endosamara racemosa (Roxb.) Geesink (Geesink 1984: 93). Geesink recognised that the plant had some unusual characters ( Table 4) that separated it from the genera in which it had previously been placed, notably that each of the 4 to 5 ellipsoid seeds is covered in a wing-like papery endocarp forming samaroid loments that enables the wind dispersal of each seed (see Plate 1B). The condition where a thin inner membranous layer surrounds the seeds forming loments is very rare among legume genera but is found in some species of the genus Sesbania Adans., Entada Adans. and in Plathymenia Benth. (Lavin and Schrire 2005: 452;Kirkbride et al. 2003: 332).
Chloroplast DNA data from the rbcL gene (Hu and Chang 2003), showed that Endosamara racemosa was sister to Millettia japonica (Wisteriopsis japonica) with BS (82%). Previously, three species were recognised in the genus; A. mahidoliae (Plate 2A, E) from Kanchanaburi Province, west Thailand; A. sericea (Plate 2B) from north-east Thailand and A. filipes (Plate 2H, I). The latter is a much more robust climber from southern China, Laos, Myanmar, Thailand and Vietnam with panicles bearing robust and thickened inflorescence axes and fragrant pale to dark bluish-lilac flowers enclosed by caducous, broad, floral bracts. This species is fully discussed above and recombined in Padbruggea. The range of both A. mahidoliae and A. sericea is now known to extend into Laos and Vietnam in regions bordering Thailand (Lôc and Vidal 2001: 13, 14).
Afgekia (without A. filipes) has several distinguishing generic characters (Tables 4, 5): stipules 10-25 mm long (the longest by far in the Callerya group); racemes axillary or terminal (panicles in A. filipes); odourless flowers (fragrant in A. filipes); callosities in two pairs on the standard petal, (a unique character in the Callerya group); stamens with a distinctive ring of retrorse hairs on the filament immediately below the anthers and seeds with hila 15-30 mm long (Table 5).
In their analysis of rbcL sequence data, Hu and Chang (2003) found Afgekia sericea placed sister to the two Wisteria species sampled. Chloroplast matK sequence data of nine species of Callerya s.l., Afgekia sericea and 11 samples from four taxa of Wisteria (Li et al. 2014) revealed, however, that Afgekia sericea was placed sister to a clade comprising Callerya s.str. This clade in turn was sister to another containing all the Wisteria samples and the two Australasian species Callerya megasperma and C. australis. Afgekia filipes was not included in this analysis. In the same paper, Li et al. (2014) also published their results from analyses of the nuclear DNA ITS spacer region. Those results showed that 14 samples of five taxa of Wisteria formed a discrete clade sister to one containing Afgekia filipes and Callerya dasyphylla sister to C. australis and C. megasperma. Afgekia sericea, however, was placed in a separate clade (with poor support) sister to Callerya eurybotrya and C. reticulata. In a majority consensus tree of the combined nuclear and chloroplast data Afgekia sericea is sister to Callerya eurybotrya and C. reticulata while Afgekia filipes is sister to Callerya australis and C. megasperma (Li et al. 2014).

Wisteria Nutt., Gen. Amer. Pl. 2: 115 (1818)
The genus Wisteria forms a distinct group of three species occurring in China, Japan and Korea and one, Wisteria frutescens, in the eastern USA. The latter is the most distinct on account of its later summer (vs. spring) flowering; standard petals reflexing near the middle vs. at the base in the Asian species; callosities of the ridge (vs. papillate) type, broad wing petals which arch above the keel with the tips adherent to each other enclosing the keel and covering the staminal column prior to anthesis vs. adherent to the keel and not as above, and the straight, non-septate, externally smooth pods containing reniform seeds vs. subseptate, velutinous, gently torulose pods containing lenticular seeds (Table 4, Plate 3F).
The analyses using plastid matK and nuclear ITS sequence data discussed above under Afgekia (Hu et al. 2002, Li et al. 2014, are the only DNA based studies to have sampled all four species of the genus Wisteria. Hu et al. (2002) included nine species of Callerya and Afgekia filipes while Li et al. (2014) included 15 species of the Callerya group. The other genera within the Callerya group, however, i.e. Endosamara, Sarcodum and Wisteria/Millettia japonica were excluded in these analyses.

Callerya group taxonomy 3: New genera within the Callerya group as delimited in this study
Our research has confirmed the uniqueness of other taxa within the Callerya group (Tables 1, 2). Schot noted the affinities between Endosamara racemosa and Callerya kityana in her revision of Callerya (Schot 1994: 25). Our results confirm that Callerya kityana with its sigmoid wing petals, among other unique autapomorphies, belongs in our new monospecific genus Sigmoidala (Fig. 3, Plate 1C-D). Our results have also revealed that two recently described Thai species, Callerya chlorantha and C. tenasserimensis (Sirichimorn et al. 2016) unequivocally share a suite of synapomorphies (Table  4) that segregate them from Callerya s.str. and they belong together in our new genus Kanburia (Plate 1E-F).
Dunn also recognised the distinctiveness of the three Australasian species M. australis, M. megasperma and M. pilipes which comprised his Sect. Austromillettia (Dunn 1912a: 140). Our results confirm Dunn's recognition and that these all belong in our new genus Austrocallerya (Fig. 6 and distinguishing characters, Table 4).
Based on a sampling of the morphologically most distinctive and apparently isolated taxon Callerya strobilifera Schot (which has not been sampled before in previous analyses), this species is placed here in the new genus Serawaia. Dunn (1912a: 135, 139) Dunn. Schot (1994) transferred all these species (except M. japonica) into Callerya and they all form a strongly supported clade in our analyses underpinned by shared morphological synapomorphies (Fig. 1, Table 4). Two of these species Callerya fordii and C. speciosa share gibbosities and glabrous standards with the other species in Dunn's Sect. Eurybotryeae but they differ in their densely pubescent ovaries and larger flowers. These two species are recognised here in the new genus Nanhaia (Fig. 4).
Geesink recognised the distinctive nature of Millettia japonica [Wisteria japonica] when he included it in a separate couplet in his key to the genera of Millettieae (Geesink 1984: 72). Millettia japonica is the only species that has not formally been treated as belonging to the Callerya group although it has been included in various molecular analyses as either Millettia japonica or Wisteria japonica (Doyle et al. 1997(Doyle et al. , 2000Kajita et al. 2001;Hu and Chang 2003). This is now the type species of our new genus Wisteriopsis which comprises the remaining species from Dunn's Sect. Eurybotreae (Fig. 5, Plate 3A-E).
Wisteria japonica Siebold & Zucc. has been recognised as distinct since the early days of European interest in Japanese botany (Fig. 5, Plate 3A-D). It was known to the physician Englebert Kaempfer who made a note of the species in the late 17 th century (Compton and Lack 2012). It was first validly described by Philipp von Siebold and Gerhard Zuccarini in their illustrated work Flora Japonica (Siebold and Zuccarini 1839: 88) based on material Siebold had collected near Nagasaki prior to 1829 (Compton and Thijsse 2013).
The American botanist Asa Gray recombined Wisteria japonica into Millettia (Gray 1858: 386) with no additional descriptive information other than "found on Kiu-siu belongs to a more southern Asiatic type" and "this is truly a Millettia". Dunn, however, in his revision of Millettia stated that there were several characters that allied the species with Wisteria, notably the deciduous, pinnate leaves and presence of what he considered to be large paniculate inflorescences of spreading to pendulous axillary racemes (Dunn 1912a: 153). Dunn also noted that there were other characters that separated the species from Wisteria, the occurrence of truly paniculate inflorescences, persistent bracts, and he asserted that the stamens of M. japonica were monadelphous (Dunn 1912a: 153). This taxon has also been accepted as belonging in Millettia in floristic works (Ohwi 1984) and in the horticultural literature (Valder 1995) and was recently maintained in Millettia in a genetic marker study (Kim et al. 2013). Geesink (1984) placed Millettia japonica close to his south-east Asian monotypic genus Endosamara racemosa in a couplet in his key to Tribe Millettieae based on their sharing paniculate inflorescences and what he perceived to have been an absence of callosities on the standard petal (see Table 4). Geesink also recognised that these two differed from each other by the uniquely compartmented and winged fruits in Endosamara and the presence of persistent bracteoles in M. japonica. He also linked M. japonica to his descriptions of Wisteria, Callerya and Sarcodum (Geesink 1984: 72, 93, 122). He considered M. japonica to be closest to, or included within, Callerya and specifically noted "the wings free from the keel in Millettia japonica" (Geesink 1984: 83), a character that he also noted for Wisteria (Geesink 1984: 121). In addition he recognised similarities between M. japonica and Sarcodum (Geesink 1984: 117) stating: Sarcodum resembles Millettia japonica (which I consider to belong to Callerya) in its habit, flower characters, and in the fleshy exocarp, but in M. japonica the pod is flat and not so convex around the seeds and this species has a "true" panicle.  in her revision of the genus Callerya, excluded Millettia (Wisteria) japonica on the basis that she believed the species had slender axillary racemes rather than panicles with thickened axes and stated that she considered the species to be closest to Callerya reticulata (Schot 1994: 5). Moreover, Schot quoted Dunn (1912a: 153) in assuming that the stamens were monadelphous. This was clearly an oversight in both cases as both Callerya (sensu  and Millettia japonica have diadelphous stamens. Schot opined that the species "lacks the facies of a 'true' Callerya" but without further comment. Our results and observations have revealed that Callerya reticulata is indeed closely related to Wisteriopsis japonica and also belongs in Wisteriopsis (Plate 3E). In addition, our studies support the inclusion of Callerya championii, C. eurybotrya and C. kiangsiensis within Wisteriopsis.
Recent molecular phylogenies that included Wisteriopsis japonica (usually as Millettia japonica) in their analyses all used data from the chloroplast gene rbcL (Doyle et al. 1997(Doyle et al. , 2000Kajita et al. 2001;Hu and Chang 2003). Doyle et al. (1997) found that a single species of Wisteria (Wisteria sp.) and Millettia japonica were sister to each other. Doyle et al. (2000) provided data in a larger dataset combined with morphology and found that two samples of Wisteria, one Afgekia and Millettia japonica formed an unresolved clade separate from the rest of Millettia. Results from a parsimony analysis by Kajita et al. (2001) which included two species of Wisteria (W. sinensis and W. sp.), Afgekia sericea and Millettia japonica found that in their strict consensus tree, the two Wisteria samples, Millettia japonica and Afgekia sericea were unresolved in a separate (but unnamed) IRLC. Hu and Chang (2003) placed Millettia japonica sister to Endosamara racemosa but these were unresolved with respect to two sister samples of Wisteria, W. sinensis and W. sp.
Wisteriopsis japonica (sampled as Wisteria japonica) has also been found to possess a unique terminal N-Acetylgalactosamine leguminous lectin which has been recognised to be useful as a probe for human lung squamous cell carcinoma (Soga et al. 2013). The molecular weight of the lectin in Wisteriopsis japonica which does not bind to galactose, is different to the molecular weights of the lectins of both Wisteria floribunda and W. brachybotrys and has a different sugar-binding specificity to the lectins of W. floribunda, W. brachybotrys and W. sinensis which all bind to galactose (Soga et al. 2013).

Integrating our results: the Callerya group to Tribe Wisterieae
In the most recent family-wide phylogenies of Leguminosae (Wink 2013;LPWG 2013LPWG , 2017, the IRLC is strongly supported, with the Callerya group and Glycyrrhiza placed in equivocal positions relative to the Temperate Tribe block). In the Maximum Likelihood (ML) tree of the most comprehensive phylogeny to date (LPWG 2017), Glycyrrhiza is sister to Schefflerodendron (outgroup used to root our analyses) and these are sister to a clade comprising the Callerya group + the Temperate Tribe block. In their Bayesian Inference (BI) analysis (LPWG 2017), the Callerya group and Glycyrrhiza are unresolved in a polytomy along with the Temperate Tribe block. Adinobotrys was not included in these analyses. In our combined analyses (Fig. 1, Suppl. material 1: Figs S1-S6, Table 3), Adinobotrys is placed without support as part of a grade sister to Glycyrrhiza + the rest of the Callerya group and then Glycyrrhiza, again without support, is sister to the remainder of the Callerya group. In the plastid analyses ( Table  3) the positions of Adinobotrys and Glycyrrhiza are switched about in the grade and in the BI analysis, Adinobotrys + Glycyrrhiza are sister to the rest of the Callerya group, all without support. In the ITS analyses, again without support, Glycyrrhiza is sister to the entire IRLC with Adinobotrys placed sister to the remainder of the IRLC excluding the rest of the Callerya group (Table 3). The equivocal positions of Adinobotrys, the residual Callerya group and Glycyrrhiza, in all recent phylogenies, points to them being relatively isolated elements compared to the Temperate Tribe block and an understanding of the relationships between them needs further research.
As discussed earlier, Jansen et al. (2008) noted that within the IRLC, the rps12 intron was uniquely present in the Callerya group but absent in all other IRLC taxa sampled, including Glycyrrhiza. Genera in the Callerya group, that they confirmed as having the rps12 intron, included species of Adinobotrys, Endosamara, Wisteriopsis, Afgekia, Padbruggea, Austrocallerya and Wisteria. Although Adinobotrys is supported as belonging within the Callerya group by Jansen et al. (2008) -and that superficially A. atropurpureus and A. vastus are very similar to other tropical genera within the Callerya group -we have decided to recircumscribe our emended Tribe Wisterieae to include the crown node of clades A to E (Fig 1, Suppl. material 1: Figs S1-S6, Tables 3, 4). This node is fully supported in both the combined and plastid analyses and in the ITS BI analysis, although only weakly so in the ITS ML analysis (Fig. 1, Table 3) and is diagnosed by the significant morphological synapomorphy of the lianescent habit. Adinobotrys may well belong within Tribe Wisterieae based on the evidence of Jansen et al. (2008), but on the basis of the consistent lack of support for its inclusion in our analyses, it is excluded here. Hu et al. (2000) also faced this quandary in that while the Phytochrome gene data (Lavin et al. 1998) and pollen data (Zhu 1994) supported linking Callerya atropurpurea with the Callerya group, their ITS and trnK/ matK evidence suggested that Callerya atropurpurea was a genetic outlier with respect to Callerya. Nevertheless, Adinobotrys is treated in full in our taxonomic treatment since it is clearly a disparate element within Callerya s.l. (sensu Schot 1994) and thus, with BS (100%) support, requires to be reinstated as a separate genus. The tree habit in Adinobotrys compared to lianas and scandent shrubs found without exception in Tribe Wisterieae serves to segregate this genus morphologically. Tribe Wisterieae together with Adinobotrys and Glycyrrhiza and the Temperate Tribe block thus represent the four main clades of the IRLC.
Tribe Wisterieae comprises a grade of three major clades (Fig. 1, Suppl. material 1: Figs S1-S6): the first branching Clade A (Sarcodum to Sigmoidala), sister to Clades B to E (Nanhaia to Wisteria) and then, Clade B (Nanhaia -Wisteriopsis) sister to Clades C-E (Callerya -Wisteria). In Clade A, the small genus Sarcodum and the monotypic Endosamara and Sigmoidala are all morphologically very distinct from each other. Sarcodum has botuliform, somewhat fleshy, pods with leaves comprising the smallest and usually most numerous leaflets in the tribe, Endosamara has unique winged and lomented seeds and Sigmoidala has distinctive 'S' shaped wing petals. Sigmoidala (as Callerya) kityana was treated as part of Callerya s.l. by  and is described here as a new genus. All three genera are fully supported at the generic level in the combined, plastid and ITS analyses as are all three together within Clade A except for the ITS ML and BI analyses where, with no support, the long-branched Endosamara appears attracted to other long branched genera within Clade C (Fig. 1, Table 3). The three genera together with Wisteriopsis (in Clade B) share glabrous ovaries as a unique character in the tribe; Sarcodum and Endosamara both share glabrous standards whereas the back of the standard in Sigmoidala is rufous pubescent and Sarcodum is the only one of the three with erect leafy racemes as opposed to the terminal panicles of the other two genera (Table 4). The genus Sarcodum has not been included in previous molecular phylogenies and is shown here to be fully supported as sister in the combined analyses to Endosamara and Sigmoidala.
Clade B (Fig. 1) comprises two new genera, Nanhaia and Wisteriopsis that both produce prominent gibbosities near the point of attachment of the leaf pulvinus to the stem (a character only shared otherwise by Serawaia in Clade C). Another synapomorphy shared by Nanhaia and Wisteriopsis is the presence of an annulus of hairs surrounding the calyx rim. Both genera also share glabrous standards with Endosamara and Sarcodum. Nanhaia and Wisteriopsis comprise species included within Callerya s.l. by , but a number of additional species in Wisteriopsis came to light as unsuspected taxa associated previously with Millettia, Wisteria and Chinese Callerya s.l. Both Nanhaia and Wisteriopsis are fully or very strongly supported as genera in all analyses except where Nanhaia is represented as a single accession (Nanhaia speciosa) in the ITS analyses ( Fig. 1, Table 3) and both are fully to strongly supported together as Clade B. Moderate support values for the sister group relationship between Clade B and Clades C-E reflect the unstable position of Clade B relative to Clade C in the ITS analyses (Table 3).
Clade C (Fig. 1) comprises the genera Callerya s.str. (Clade C1), sister to a moderately to strongly supported alliance (in the combined analyses) of Clades C2 + C3 + C4 (Table 3), comprising Whitfordiodendron, Serawaia, Kanburia and Afgekia This grouping is poorly supported in the ITS analyses, however, and it breaks apart somewhat in the plastid analyses. Clade C as a whole is moderately (BS) to strongly (BPP) supported in the combined analyses but tends to break up and group with other genera in the plastid and ITS results ( Table 3). The genera Afgekia, Kanburia and Whitfordiodendron are each fully supported, as is Callerya s.str. at the node above C. bonatiana. Callerya s.str., including C. bonatiana, is well supported in the combined and plastid BI analyses (Table 3) but only poorly so in the ML analyses. The ITS results either split Callerya s.str., placing C. bonatiana in a polytomy with the long-branched Endosamara (from Clade A2) in the ML analysis, or in an unsupported position sister to Clade B in the BI analysis. Callerya bonatiana shares the morphological synapomorphies of Callerya s.str., e.g. wings shorter than the keel, and none of the synapomorphies of other Wisterieae genera so it is treated here as part of Callerya s.str. The genus Whitfordiodendron is reinstated at generic level from Callerya s.l. ) and a new genus Kanburia is described from only recently collected material (Sirichamorn et al. 2016). Sister group relationships with Kanburia (Clade C3) are generally not supported, although in the ITS analyses the strongest (although still poor) support is with Serawaia (Table 3). In the combined BI analysis, however, there is strong support (0.95 BPP) for Afgekia being sister to Kanburia. The genus Serawaia is the only new genus described here that is based on a single accession since the DNA of all other material sampled was too degraded to be useful. It is a morphologically unique taxon in the Tribe Wisterieae with large cone-like strobilate inflorescences of bright yellow flowers with the persistent 15-18 × 8-12 mm bracts becoming indurate and coriaceous in fruit. No other taxon has this combination of characters and while it is placed sister to Whitfordiodendron with no support in the combined analyses and sister to Kanburia with poor support in the ITS analyses ( Fig.  1, Table 3), it remains strongly supported within Clade C (BS 87%, BPP 0.98) in the combined analyses. At the same time Serawaia cannot be placed morphologically with any other genus in Clade C. Morphologically, the presence of gibbosities below the stipules and wing petals free of the keel are unique in Clade C but are characteristic of all or some taxa in Clade B. No support is found, however, for links between Serawaia and clade B in these analyses.
Afgekia comprises two very distinct species separated from the other genera in the clade by a long branch and a large number of synapomorphies, most notable of which are the racemes vs. panicles present in the rest of Clade C, long sericeous floral bracts and, uniquely within the tribe, the two pairs of callosities on the standard petals (Table 4). The seeds of Afgekia have long strap-shaped hila compared to the short elliptic hila in the other genera of Clade C. Callerya is distinguished by having wing petals much shorter than the keel, Whitfordiodendron by its densely sericeous keels (glabrous elsewhere in the tribe although pubescent to tomentose in Afgekia) and Afgekia and Kanburia both lack bracteoles (present in Callerya, Serawaia and Whit-fordiodendron). Afgekia, Callerya, Kanburia, Serawaia and Whitfordiodendron all share densely pubescent or sericeous ovaries and backs to the standard petals. The pods of Whitfordiodendron are inflated, ovoid, rugose to ridged or ruminate (Fig. 2P), with 1-2(-3) seeds becoming fused together when more than one in the pod. The pods in Afgekia are inflated with densely velutinous surfaces (vs. in Callerya, Serawaia and Kanburia pods are flattened to inflated, smooth, with (1-)2-6 seeds remaining separate in the pod (Fig. 2L).
The genera Padbruggea and Austrocallerya form a strongly supported Clade D (Table 3) in the combined, plastid BI and ITS analyses although support is weak in the plastid ML analysis. Padbruggea and Austrocallerya are also strongly supported as genera in all analyses. Padbruggea is reinstated in part from Callerya s.l.  and by the transfer of P. filipes from Afgekia. Austrocallerya is described as a new genus comprising the Australasian species in Callerya s.l. . The two genera share seeds with long, strap-shaped hila and more open, laxly flowered panicles as morphological synapomorphies and are segregated from each other based on standard callosity shape, fruit characters and geographic distribution. Clade D is sister to Clade E (comprising Wisteria) with strong support in all results (Table 3) except in the plastid ML analysis.
Four species of Wisteria, three in temperate east Asia (Clade E1) and one in North America (the only non Asian-Australasian species in the Wisterieae, Clade E2) comprise Clade E with full support in the combined, plastid and ITS analyses (Table 3). Clade E is distinguished from Clade D by its deciduous vs. evergreen habit, racemose vs. paniculate inflorescences, seeds up to 12 mm in size in compressed pods vs. seeds larger than 12 mm in inflated pods and seeds with short elliptic hila vs. long strap-shaped hila ( Table 4). The deciduous habit is only found otherwise in Wisteriopsis japonica.
Aberrant results in the ITS analyses compared to those derived from plastid data are thought likely to be the outcome of fewer representative taxa, probable long branch attraction in the placement of Endosamara, Kanburia, Serawaia and Afgekia and alignment problems with ITS making it difficult to ascertain true homology. The 77.1% pairwise percentage of identity across the ITS alignments vs. 92.5% for the overall plastid alignments may also be indicative of ITS being less informative than the plastid data in these analyses. It is also apparent that the two single accessions of Callerya bonatiana and Serawaia strobilifera (both limited by a lack of research material) are the most labile in the phylogeny, thereby reducing the support values of their associated clades.

Taxonomic treatment of Tribe Wisterieae
Thirteen genera within a much expanded Tribe Wisterieae are described here, encompassing five clades recovered in our phylogenetic analyses (Fig. 1). In addition, the genus Adinobotrys is fully described owing, in part, to its long association with the Callerya group but also because of its equivocal sister group relationships to tribe Wisterieae in these analyses. The following clades and included genera are listed below (Fig. 1 Note. The Tribe Wisterieae is distinguished by comprising woody lianas or sprawling scandent shrubs. All species have bracts that in the most part enclose immature buds at the apex of inflorescences and all bear either true panicles or true racemes as opposed to pseudopanicles and pseudoracemes. The tribe is further distinguished from Tribe Millettieae by all genera lacking one 25 kb long copy of the inverted repeat in the chloroplast genome. Morphological key to the genera in Tribe Wisterieae together with Adinobotrys Large spreading evergreen trees to 20 m or more in height. Stems green when young, terete, finely brown pubescent becoming brown and glabrous with age. Leaves with 5-9 (-11) leaflets, evergreen, coriaceous and nitent when mature, imparipinnate, rachis 11-33 cm long. Stipules 2-4 mm long, deltoid, persistent. Stipels absent. Leaflets 5-21 × 2-11 cm, ovate, elliptic or obovate, glabrous above and below, upper surface nited, apex acute to acuminate, margins entire, base obtuse or cordate. Inflorescence a robust many-flowered erect terminal panicle 10-40 cm long, peduncle sparsely hairy to tomentose. Flowers 14-20 mm long, emerging from February to May (in A. atropurpureus) and May to November (in A. vastus). Floral bracts 2-4 mm long, persistent (caducous in A. vastus), ovate. Bracteoles 1-2 mm long, at base of calyx tube, persistent, ovate. Pedicels 2-6 mm long, densely pubescent. Calyx narrowly campanulate, oblique, green, tube 4-6 × 6 mm, puberulent externally, five lobed, lobes unequal 0.5 mm long, acute or obtuse. Standard 11-20 × 13-20 mm broadly ovate, inner surface pink, dark reddish-purple, rarely white, nectar guide yellow, back of standard glabrous, apex acute, callosities of boss type. Wing petals 12-19 × 5-8 mm, glabrous, ± equal or longer than keel in length, each broadly semipandurate with basal claws 3-5 mm long. Keel petals 12-18 × 9 mm, glabrous, apex acute to rounded. Stamens diadelphous, nine fused together, the vexillary one free, all curved upwards at apex. Ovary sparsely to densely hairy, style glabrous, 5-6 mm long curved upwards at apex, stigma punctate. Pods 7-25 × 3-6 cm, inflated or flat (A. vastus), irregularly ovate to oblong or narrowly elliptic, dehiscent, surface glabrous, finely rugose, subseptate. Seeds 1-4, irregularly ovoid to oblong or flattened orbicular, sometimes laterally compressed inside the pod, 15-38 × 20-35 × 3-26 mm, hilum 2-3 × 2 mm, ovate-elliptic or circular.

Key to species of Sarcodum
Habitat. In dry woods, thickets and forest margins from sea level to 850 m. climbing over rocks, on banks and among scrub and trees. Diagnosis. The monospecific Sigmoidala kityana has several affinities with Endosamara racemosa including the absence of bracteoles and glabrous ovaries, characters which also separate it from Callerya s.str. which has bracteoles and sericeous ovaries. Sigmoidala also shares with Endosamara the pubescent floral bracts and broadly campanulate, slightly oblique, subtruncate calyx as noted by Schot (1994: 25) but it was placed in Callerya on account of the fruits that lacked the lomented endocarp. The stipules in Sigmoidala are shorter, 3-7 mm long (vs. 9-12 mm in Endosamara); pedicels shorter, 3-4 mm long (vs. 4-12 mm in Endosamara and Sarcodum), floral bracts linear, 6-8 mm long (vs. linear-lanceolate, 8-12 mm long in Endosamara, 6-20 mm in Sarcodum); the back of the standard densely, appressed rufous pubescent (vs. glabrous in Endosamara and Sarcodum) and the wing petals of Sigmoidala are unique within Tribe Wisterieae being a sigmoid shape, reflexed at the midpoint and extending outwards towards the apex (see Fig. 3I). The pods of Sigmoidala are flattened, linear to obovate, 7-11 × 1-2 cm (vs. septate, flattened, linear, 10-25 × 1-2 cm in Endosamara and botuliform in Sarcodum).
Key to species of Callerya recognized in this treatment  Illustrations. Lôc  Diagnosis. This monospecific genus has several autapomorphies compared with other genera within the tribe. It is the only species that has large and very persistent imbricate floral bracts along the inflorescence enclosing the uniquely golden-yellow flowers. Serawaia is the only genus in Clade D that has prominent gibbosities below the stipules. Its nearest affinities lie with Callerya, Kanburia and Whitfordiodendron which all have sericeous backs to their standard petals. The back of the standards of Serawaia are, however, pubescent but the hairs are not as long as those in Afgekia, the other member of Clade D. The wing petals, which are free from the keel, almost equal the length of the keel as in Kanburia, Whitfordiodendron and Afgekia which distinguishes these four genera readily from Callerya whose wings are shorter. The ovary in Serawaia is only sparsely hairy whereas in all four other genera within Clade D the ovaries are densely sericeous (see figs 4 and 5 in Schot (1994: 33, 34).
Etymology. named after the Serawai river in west Kalimantan, a tributary of the Kapuas river, where the species was first discovered. Habitat. In open sites climbing among trees and scrub on exposed ridges and riverbanks from sea level to 350 m.  fordiodendron (vs. glabrous in Kanburia and Callerya). The pods in Whitfordiodendron are inflated and ovoid with a velutinous or pubescent surface (vs. linear, compressed, glabrescent in Kanburia). The ovoid seeds in Whitfordiodendron may become fused together when there are more than one per pod (vs. lenticular, separate in pod in Kanburia). The wing petals are equal in length with the keel petals in Whitfordiodendron (vs. shorter in Callerya).

Key to species of Padbruggea
Illustration. Lectotype sheet of Padbruggea dasyphylla at (L); L1978535. Distribution. Indonesia (Borneo: Kalimantan, Java, Sumatra); Malaysia (Peninsula, Sarawak); Thailand.  (1891) Nomenclatural note. There is a sheet at Kew, K000881019, with two different collections by Maingay. One has two mature pods, a leaflet and a few scraps of stem, the other has several leaflets and bits of stem. There are two labels attached at the bottom of the sheet; one states "Herbarium A.C.Maingay 2757, Singapore, 1867-1868, apparently a climber, no duplicates of this interesting sp.". The other has "Herbarium of the late A.C.Maingay 605, Malaya, distributed at the Royal Gardens, Kew, 1871". There is, however, no indication as to which collection represents Maingay 2757 and which might be Maingay 605. Baker in his protologue mentions "Singapore, Maingay" and described the 15 or more leaflets, rounded at both ends and the oblong, velvety pod traversed with deep longitudinal grooves. Since Baker described both fruit and leaves we have inferred that the left-hand fruiting specimen is Maingay 2757 and thereby have selected it to lectotypify the name. Dunn (1911: 197) in his key to the species Padbruggea dasyphylla and P. maingayi stated that P. dasyphylla had leaflets with revolute margins and was densely tomentose below whereas P. maingayi did not have leaflets with revolute margins and was nearly glabrous below. Our examination of type material of both species has found that the reverse is the case as indicated in our key to the species.
Distribution. Indonesia (Java); Malaysia (Peninsula); Singapore. ( Note. Dunn (1912a) recognised the distinctiveness of the Australasian species when he placed all three in his Millettia Sect. Austromillettia Dunn. He noted the single flowers as opposed to flowers in pairs (sometimes more than two branching from the same place on the inflorescence axis in other Millettia spp.), and the terete woody nature of the pods (Dunn 1912a: 135, 138, 140).

Diagnosis.
Austrocallerya comprises three Australasian species with glabrous or finely pubescent young leaves and stems (vs. these densely brown tomentose in Padbruggea, see Table 4 and Fig. 6). The robust paniculate inflorescences are more erect than those in Padbruggea and the flowers have very broad standard petals with a recessed dividing midline. Either side of the midline is an arch callosity which forms a short crescent arching over the staminal sheath (vs. papillate or ridge callosities in Padbruggea). The pods are fusiform (vs. obovoid or compressed-cuboid in Padbruggea), torulose and with either longitudinal striations and furrows (A. megasperma), or with irregular fine striations (A. australis) or smooth (A. pilipes), the surface in all cases being densely velutinous or pubescent. The pods of Austrocallerya can be distinguished from those of Padbruggea, which are also densely velutinous, by their outline. Padbruggea pods are either obovoid (in P. filipes) or oblong with a prominent dorsal midline flanked by two large flanges meeting at the apex (P. dasyphylla). The pods in Austrocallerya are 30-52 mm wide (vs. 40-110 mm wide in Padbruggea). The 2-10 seeds in Austrocallerya are oblong, ellipsoid or globose, frequently with one side compressed within the pod (vs. 1-2 elliptic-ovoid or prolatespheroid seeds which may also be laterally compressed in Padbruggea). In Austrocallerya the strap-shaped hila are 16-30 × 2-4 mm, (vs 16-40 × 5-10 mm in Padbruggea). Fig. 6.
Distribution. Australia (New South Wales, Queensland); Papua New Guinea (Bougainville Island, New Britain Island); New Caledonia; Cook Islands.
Habitat. In rainforest or in dry forest from sea level to 1600 m, climbing up trees and over shrubs.
Etymology. The generic name reflects the southern hemisphere distribution of the genus, austro -"australis" = south (Latin) and "callerya" a reference to their former generic placement and affinity.  We have chosen MEL2144485 collected by Mueller and Hill from the Pine River near Moreton Bay as cited in the protologue as lectotype. 2) Wisteria involuta Sprague (Sprague 1904: 141) was described from cultivated material at K. The material was collected from the Richmond River area of NSW, Australia, collector unknown. Sprague (1905: 3) recombined the species as Derris involuta (Sprague) Sprague the following year having seen the flat, one-seeded, winged fruits of the millettioid genus Derris Lour.  Sprague, Gard. Chron. Ser. 3., 36: 141 (1904) Note. There is also a specimen at K collected by Frederick M. Bailey from the Johnstone River in August 1892 -K000880982. Bailey, the author of the name, had been Colonial Botanist for Queensland since 1881 but Schot's choice of this specimen as an isotype (Schot 1994: 29) is incorrect as the holotype cited in the protologue was collected by Thomas Lane Bancroft, moreover, Bailey's specimen was collected two years after the protologue was published.
Habitat. In rainforest climbing trees and over scrub from 300 to 1200 m.
Habitat. In temperate forests from sea level to 1800 m, climbing among trees and shrubs.
Etymology. The generic name commemorates the anatomist Professor Caspar Wistar (1761-1818), President of the American Philosphical Society. It also commemorates Caspar Wistar's cousin Charles Jones Wister (1782-1865), friend of Thomas Nuttal who was the author of the name. Habitat. In clearings of evergreen lowland forest and along riverbanks at sea level to 650 m.

Key to species of Wisteria
Note. Plants from more southerly regions either side of the Appalachian mountains have previously been recognised as a separate species Wisteria macrostachya (Nutt. ex Torr. & A.Gray) B.L.Rob. & Fernald, Gray Man. Bot. N. U.S. ed. 7: 515. 1908. Observation of living plants and careful examination of many herbarium specimens coupled with the DNA generated results from this study have led us to conclude that there is only the single species W. frutescens representing the genus Wisteria in North America. We also conclude that there is sufficient difference between the southern plants and those from further north to recognise the southern one at the rank of subspecies. At the rank of species the name Diplonyx elegans Raf. (1817: 101) has priority over the widely used name Wisteria macrostachya (Robinson and Fernald 1908: 515), however, the combination Wisteria elegans has never been made (for a more comprehensive discussion see Compton (2015a)). Plants from the northerly range of the species (subsp. frutescens) have smaller and shorter inflorescences without (or with very few) glandular hairs on pedicels and calyces (vs pedicels and calyces covered in clavate glandular hairs in subsp. macrostachya). The teeth on the calyces of subsp. frutescens are ± subequal (vs lower teeth much longer in subsp. macrostachya). Racemes from the colder north (subsp. frutescens) are usually considerably shorter than the elongating racemes of southerly plants (subsp. macrostachya). For a more comprehensive description of these taxa see Compton and Lane (2019 Habitat. Climbing over trees and shrubs and along banks and over thickets 50 to 1800 m.