Transfer of Millettia pachycarpa and M. entadoides to Derris (Fabaceae), supported by morphological and molecular data

Abstract

Derris Lour. and Millettia Wight & Arn. are two genera of the tribe Millettieae (Fabaceae). The biggest difference between the two genera is that Derris has indehiscent, thin and winged pods, while Millettia has dehiscent, compressed and wingless pods, but some exceptions also exist. Because of their tardily dehiscent, inflated and wingless pods, two Asian liana species, Millettia pachycarpa Benth. and M. entadoides Z. Wei, were previously placed in Millettia sect. Macrospermae Dunn. However, their vegetative and floral characters, as well as the shape of seeds match with those of Derris very well. Phylogenetic analyses based on three chloroplast markers (trnK/matK, trnL-F IGS, and psbA-trnH IGS), and nuclear ITS/5.8S showed that the two species are nested within Derris, and separate from other samples of Millettia species with strong support. Therefore, the two species should be transferred from Millettia to Derris to maintain the monophyly of the latter, which inevitably results in an emendation of the generic circumscription of Derris. Due to the existence of Derris pachycarpa Merr., the second earliest valid epithet “taiwaniana” should be used for this taxon when transfering Millettia pachycarpa Benth. to Derris. Thus, a new combination, Derris taiwaniana (Hayata) Z.Q. Song, is proposed here. Millettia entadoides Z. Wei is recombined as Derris entadoides (Z. Wei) Z.Q. Song. The floral morphology of D. entadoides is described here for the first time, and it is reported as a new record from Thailand in this study.

Full text

Phytotaxa 531 (3): 230–248
https://www.mapress.com/pt/
Copyright © 2022 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
230 Accepted by Yi-Hua Tong: 8 Jan. 2022; published: 21 Jan. 2022
https://doi.org/10.11646/phytotaxa.531.3.4
Transfer of Millettia pachycarpa and M. entadoides to Derris (Fabaceae), supported
by morphological and molecular data
ZHUQIU SONG1, 3* & BO PAN2, 4
1 Key Laboratory of Plant Resources Conservation and Sustainable Utilization, South China Botanical Garden, Chinese Academy of
Sciences, Guangzhou 510650, P. R. China.
2 Center for Integrative Conservation, Core Botanical Gardens, Xishuangbanna Tropical Botanical Garden, Chinese Academy of
Sciences, Menglun, Mengla 666303, P. R. China.
3
�
songzhuqiu@scbg.ac.cn; https://orcid.org/0000-0002-7763-2431
4
�
pb@xtbg.org.cn; https://orcid.org/0000-0002-0038-9664
*Author for correspondence:
�
songzhuqiu@scbg.ac.cn
Abstract
Derris Lour. and Millettia Wight & Arn. are two genera of the tribe Millettieae (Fabaceae). The biggest difference between
the two genera is that Derris has indehiscent, thin and winged pods, while Millettia has dehiscent, compressed and wingless
pods, but some exceptions also exist. Because of their tardily dehiscent, inflated and wingless pods, two Asian liana
species, Millettia pachycarpa Benth. and M. entadoides Z. Wei, were previously placed in Millettia sect. Macrospermae
Dunn. However, their vegetative and floral characters, as well as the shape of seeds match with those of Derris very well.
Phylogenetic analyses based on three chloroplast markers (trnK/matK, trnL-F IGS, and psbA-trnH IGS), and nuclear ITS/
5.8S showed that the two species are nested within Derris, and separate from other samples of Millettia species with strong
support. Therefore, the two species should be transferred from Millettia to Derris to maintain the monophyly of the latter,
which inevitably results in an emendation of the generic circumscription of Derris. Due to the existence of Derris pachycarpa
Merr., the second earliest valid epithet “taiwaniana” should be used for this taxon when transfering Millettia pachycarpa
Benth. to Derris. Thus, a new combination, Derris taiwaniana (Hayata) Z.Q. Song, is proposed here. Millettia entadoides Z.
Wei is recombined as Derris entadoides (Z. Wei) Z.Q. Song. The floral morphology of D. entadoides is described here for
the first time, and it is reported as a new record from Thailand in this study.
Keywords: Asia, Leguminosae, Millettieae, new combination, systematic position
Introduction
Derris Loureiro (1790: 432) and Millettia Wight & Arnott (1834: 263) are two large genera within the tribe Millettieae
of the family Fabaceae (Geesink 1984, Schrire 2005, Wei & Pedley 2010). Derris was recently united with Paraderris
(Miquel 1855: 145) Geesink (1984: 109), but separated from Brachypterum (Wight & Arnott 1834: 264) Bentham
(1837: 37) (Sirichamorn et al. 2012a, 2014a, b). The re-circumscribed Derris comprises about 55–60 species of woody
climbers that are distributed mostly in Southeast Asia (Sirichamorn et al. 2014c). Millettia includes 90–200 species of
woody climbers, shrubs, or trees from Asia and Africa (Song et al. 2021a, b), and phylogenetic analyses showed that it
is polyphylectic in its current circumscription (Hu et al. 2002, LPWG 2017, Cooper et al. 2019). The two genera have
very similar appearance, especially the morphological characters of inflorescences and flowers, which has resulted
in the transfer of some names between them (Merrill 1910, Gagnepain 1913, How 1954, Song et al. 2017). Derris
generally has indehiscent, thin and two-winged pods, while Millettia has dehiscent, compressed and wingless pods
(Dunn 1912, Sirichamorn et al. 2012b, 2014b, Mattapha 2017, 2020). However, exceptions exist in both genera. For
example, the pods of some Millettia species are indehiscent [e.g., Millettia pinnata (Linnaeus 1753: 741) Panigrahi in
Panigrahi & Murti (1989: 210)] or winged [e.g., M. tetraptera Kurz (1873: 69)], and some Derris species have inflated
and almost wingless pods, such as D. montana Bentham (1852: 253). Nevertheless, the fruit characters have long been
considered as the biggest difference between the two genera (Geesink 1984, Hu et al. 2000, Wei & Pedley 2010, Song
et al. 2017, Mattapha 2020).

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Two Asian species, Millettia pachycarpa Bentham (1852: 250) and M. entadoides Z. Wei (1985: 278), were
described as woody climbers with tardily dehiscent, inflated and wingless pods (Wei 1994, Sun 2006, Wei & Pedley
2010), and therefore were previously placed in Millettia sect. Macrospermae Dunn (1912: 136). But “tardily dehiscent”
pod is difficult to define, and is recognized as the same type as indehiscent pod sometimes. The type specimens of M.
entadoides, which were collected in March, have pods that remain indehiscent (Fig. 1), and we also found that the old
pods were still indehiscent even in April in the field (Fig. 2). The fruit valves of M. pachycarpa are soft and leathery,
and appear to be not separate even at maturity, as pointed out by Dunn (1912: 132) and observed by us in the field (Fig.
3). Therefore, the two species seem to be intermediate between Millettia and Derris with respect to the characters of
pods. In fact, the importance of the fruit characters, including the dehiscence of fruits and the presence or absence of
wings along sutures, have long been overestimated in the classification of Millettieae as evaluated by Hu et al. (2000)
based on molecular evidence.
Recent molecular phylogenetic analyses have revealed the close relationship of Millettia pachycarpa with Derris
rather than Millettia (LPWG 2017, Cooper et al. 2019, Hu et al. 2020). For instance, a comprehensive phylogenetic
analysis of the family Fabaceae based on the plastid matK sequences showed that M. pachycarpa is mostly related to
some species of Derris such as D. montana with inflated pods, and is separated from other species sampled in Millettia,
although the support value was relatively weak (LPWG 2017). For Millettia entadoides, it is a hitherto poorly known
species from China, for which the floral characters were never described in previous studies (Wei 1985, 1994, Sun
2006, Wei & Pedley 2010). In his PhD thesis, Mattapha (2017) described a new species of Derris and compared it
with M. entadoides in morphology, but no DNA sample of M. entadoides was sequenced. In order to investigate the
phylogenetic position of Millettia pachycarpa and M. entadoides within the tribe Millettieae, the DNA data from
genome skimming were used in this study.
Material and methods
Morphological observations
Observations of morphological characters were mainly based on herbarium specimens. We examined more than 500
collections of Millettia entadoides and M. pachycarpa kept in the herbaria A, AU, BM, CDBI, CPNP, CSFI, CSH,
CZH, E, FJSI, GFS, GNUG, GXMG, GXMI, GYBG, GZAC, GZTM, HGAS, HITBC, HN, IBK, IBSC, IMC, IMDY,
JJF, JJU, JXU, K, KUN, L, LBG, MO, NAS, NF, NY, P, PE, SM, SYS, SZG, TAI, TCD, US, WUK, XZ, and ZM.
Acronyms for the herbaria follow the Index Herbariorum (Thiers 2021). We also conducted several field investigations,
especially for the poorly known M. entadoides.
DNA sequencing and phylogenetic analyses
A total of 120 species from 26 genera, including 25 species (30 samples) of Derris and 31 species (41 samples) of
Millettia were included as ingroups in this study (Appendix 1). Three species of the genus Indigofera Linnaeus (1753:
751) were used as outgroups. The ingroups and outgroups were selected according to previous phylogenetic analyses
of Derris-like taxa within the tribe Milletieae by Sirichamorn et al. (2012a) and of the whole family Fabaceae by
LPWG (2017). Two samples of Millettia pachycarpa, one sample of M. entadoides, and one sample of Derris fordii
were newly sequenced in this study. The remaining sequences were downloaded from Genbank (www.ncbi.nlm.nih.
gov/Genbank).
Three chloroplast markers, trnK/matK, trnL-F IGS (intergenic spacer) and psbA-trnH IGS, and one nuclear
marker ITS/5.8S, were used in this study following Sirichamorn et al. (2012a). We obtained these DNA regions of
new samples using the genome skimming method (Zeng et al. 2018). We conducted paired-end (PE) sequencing on
the Illumina HiSeq X-Ten instrument at Beijing Genomics Institute (BGI) in Wuhan, China. The sequences were
assembled using GetOrganelle v1.7.5 (Jin et al. 2020), annotated using PGA program (Qu et al. 2019), and checked
using Geneious 11.0.4 (Kearse et al. 2012).
We performed sequence alignment using MAFFT v.7.450 (Katoh & Standley 2013) with LINSI algorithm in
Geneious 11.0.4. Three DNA datasets were used in this study – ITS/5.8S, combined chloroplast sequence (trnK/matK,
trnL-F IGS, and psbA-trnH IGS), and combined nuclear and chloroplast sequence, which contain 354 parsimony-
informative characters of 823 positions, 1077 of 4327, and 1431 of 5150 after alignment, respectively. We conducted
phylogenetic analyses using maximum likelihood (ML) with the IQ-TREE algorithm (Nguyen et al. 2015) on the W-
IQ-TREE web server (http://iqtree.cibiv.univie.ac.at, Trifinopoulos et al. 2016).

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FIGURE 1. Isotype of Derris entadoides (Z. Wei) Z.Q. Song (Chi-Wu Wang 72150, A00283892, https://kiki.huh.harvard.edu/databases/
specimen_search.php?barcode=A00283892). © Copyright of the Herbarium of the Arnold Arboretum of Harvard University.

MILLETTIA PACHYCARPA AND M. ENTADOIDES TO DERRIS Phytotaxa 531 (3) © 2022 Magnolia Press • 233
FIGURE 2. Derris entadoides (Z. Wei) Z.Q. Song. A. habit; B. stem with brown lenticels; C. cross section of stem, showing pithy inside;
D. leaf, adaxial view; E. leaf, abaxial view, showing glaucous surface beneath; F. flowering branchlets; G. pseudoraceme, showing distinct
brachyblasts; H. part of inflorescence with opened flowers; I. part of inflorescence with unopened flowers; J. brachyblast with one flower,
and dissection of a flower, scale bar = 1 cm; K. part of ovary, showing 5 ovules; L. old pods, scale bar = 4 cm. Photographs by Zhu-Qiu
Song

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FIGURE 3. Derris taiwaniana (Hayata) Z.Q. Song. A. habit; B. stem with brown lenticels; C. leaflets, adaxial and abaxial view, scale
bar = 2 cm; D. leaves; E. flowering branchlets; F. pseudoraceme, showing distinct brachyblasts; G. part of inflorescence; H. part of
inflorescence with unopened flowers; I. flower and dissection of a flower, scale bar = 2 cm; J. part of ovary, showing 7 ovules; K. old pods
covered with warts outside and reniform seeds, scale bar = 4 cm. Photographs by Zhu-Qiu Song (A–C, E–J), You-Sheng Chen (D), and
Bo Pan (K).

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FIGURE 4. Consensus tree of maximum likelihood (ML) analysis of the combined dataset of all four markers (trnK-matK, trnL-F IGS,
psbA-trnH IGS and ITS). Numbers near branches are SH-aLRT and ultrafast bootstrap (UFBoot) support values. Samples of Derris
species were indicated with red fonts, and those of Millettia species were indicated with blue fonts. The Paraderris subclade (i.e. the DP
subclade) within the clade D2 of the genus Derris is indicated in the shaded box.

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FIGURE 5. Distribution of Derris entadoides (Z. Wei) Z.Q. Song (red triangle) and Derris taiwaniana (Hayata) Z.Q. Song (blue dots,
made by the software ArcGIS 10.2).
Results
The ML analyses of the three DNA datasets resulted in consensus trees with very similar topologies. Thus, we only show
the consensus tree for combined nuclear and chloroplast sequence here (Fig. 4). The core Millettieae was recognized as
a monophyletic group with strong support (SH-aLRT and UFBoot is 99.5% and 100%, respectively). Two main groups
within Millettieae were further recognized, i.e., the canavanine group (99.7%, 100%) and the non-canavanine group
(100%, 100%). The 41 samples of 31 Millettia species did not group together and were clustered into six different
clades (Clade I–VI) in our analyses. Most of the six clades were strongly supported in the consensus tree (SH-aLRT >=
80%, UFBoot >= 95%). The clade VI (100%, 100%) was composed of 25 species of Derris and 2 species of Millettia
(M. pachycarpa and M. entadoides), and clustered into two subclades, i.e., subclade D1 (98.3%, 100%) and subclade
D2 (99.9%, 100%). Furthermore, three samples of M. pachycarpa were grouped together (60%, 98%) and most related
to one sample of M. entadoides (96.2%, 98%). The four samples of Millettia together nested within the Paraderris
subclade (i.e., the DP subclade in Fig. 4).
Discussion
Our phylogenetic analyses produced similar relationships among taxa in the tribe Millettieae as previous studies
(Sirichamorn et al. 2012a, 2014a, LPWG 2017). The results showed that the core Millettieae is comprised of canavanine
and non-canavanine groups, which corresponds to the species with and without canavanine in the seeds (Schrire 2005).
Millettia was a polyphyletic genus, consisting of taxa from six clades in our results, which is also confirmed by
previous analyses based on PHY (Lavin et al. 1998), trnK/matK (Hu et al. 2000, Queiroz et al. 2015, LPWG 2017),
rbcL (Kajita et al. 2001), ITS (Hu et al. 2002), or a combined dataset of several DNA markers (Cooper et al. 2019, Hu

MILLETTIA PACHYCARPA AND M. ENTADOIDES TO DERRIS Phytotaxa 531 (3) © 2022 Magnolia Press • 237
et al. 2020). Our results also agree with those of Sirichamorn et al. (2014a), in which Derris was strongly supported
to be united with Paraderris but separated from Brachypterum. However, Derris as proposed by Sirichamorn et al.
(2014a) was resolved as non-monophyletic in our results, because Millettia pachycarpa and M. entadoides were deeply
embedded within Derris. These two Millettia species were shown to be clearly separate from other species of Millettia,
but closely related to species from the Paraderris group. Therefore, in order to maintain the monophyly of Derris,
Millettia pachycarpa and M. entadoides (with tardily dehiscent and wingless pods) need to be treated as members
of Derris, which will result in an emendation of the generic circumscription of Derris. Of the two synapomorphies,
i.e. the liana habit and two-winged pods, proposed in Sirichamorn et al. (2014a), only the first remains in this re-
circumscribed Derris. Moreover, Derris then includes both indehiscent and tardily dehiscent pods.
In fact, when publishing Millettia pachycarpa, Bentham (1852) referred to its similarity with Derris ferruginea
Bentham (1852: 252). Some authors also indicated that D. elliptica Bentham (1860: 111) is very similar to M. pachycarpa
in flowers and leaves (King 1897, Brandis 1906). Examination of specimens and field observations revealed that M.
pachycarpa and M. entadoides have the habit of lianas, distinct lenticels on branchylets, leaves without stipels, several
pseudoracemes inserted on lower part of branchlets of current year, 3–5 (usually 3) flowers on the top of each distinct
brachyblast, reddish calyx, whitish to purplish petals, standard petals reflexed at the base and with a basal green-
yellowish patch, keel petals with lateral pockets, hardly visible floral disk, monadelphous stamens with basal fenestrae,
and reniform seeds with the hilum in a submedial sinus (Fig. 2–3). In these characters, both M. pachycarpa and M.
entadoides are consistent with the genus Derris, especially the species of the Paraderris subgroup (Sirichamorn et al.
2012b). In contrast, Millettia exhibits high variations in these characters, such as the habit of liana, shrubs or trees, the
leaves with or without stipels, the axillary, terminal or cauliflorous pseudoracemes, the indistinct or distinct floral disk,
and various shape (usually lens-shaped or quadrate) of seeds (Dunn 1912, Polhill 1971, Geesink 1984, Wei & Pedley
2010, Mattapha 2020). Moreover, Millettia pachycarpa contains rotenone toxin in its seeds and roots and was used as
a fish poison, which is also similar to most Derris species (Chiu 1950, Singhal et al. 1982, Sirichamorn et al. 2012a).
Therefore, based on the results of molecular and morphological analyses, Millettia pachycarpa and M. entadoides are
formally transfered to Derris, as treated below.
Taxonomic treatment
1. Derris entadoides (Z. Wei) Z.Q. Song, comb. nov.
≡ Millettia entadoides Z.Wei (1985: 278, fig. 7). Type:—CHINA. Yunnan, Chen-Kang Hsien [Zhenkang County], mountain slope, 1500
m, March 1936, Chi-Wu Wang 72150 (holotype: PE, not seen; isotypes: A00283892!, KUN0645004!, LBG00094202!).
Liana large, climbing. Stems cylindric, dark brown, densely scattered with brown lenticels, pithy inside; young shoots
yellow tomentose, glabrescent when mature. Stipules ovate-triangular, ca. 2 × 2.5 mm, rounded to acute at apex.
Leaves imparipinnate, 9–13-foliolate (usually 11), juvenile at anthesis, reddish when young; rachis 9.7–21.6 cm long,
including petiole 5.1–11.6 cm long; leaflet blades oblanceolate to oblong, 4.5–12.8 × 1.5–4.6 cm (ratio usually 2–4),
papery to subleathery, glabrous above, appressed puberulent to glabrescent and glaucous beneath, acuminate at apex,
cuneate to rounded at base; lateral veins 9–12 on each side of midvein, obvious, looped near margin; petiolules 5.5–6.5
mm long; stipels usually not persistent even on young leaves but occasionally present. Pseudoracemes 5–15 cm long,
usually inserted in the leafless axil of leaf on the lower part of branchlets of current year; rachis appressed tawny hairy;
brachyblasts distinct, wart-like or knob-like, up to 4 mm long and 1.5 mm wide on the lower part of the inflorescence,
with 3–5 flowers on the top of each brachyblast; bracts inserted at base of brachyblasts and pedicels; bracteoles 2,
inserted at the base of calyx. Flowers pink, ca. 1.3 cm long; pedicels 0.4–0.6 cm long, appressed tawny hairy; calyx
campanulate, green at first and then becoming dark red, 4-lobed, with very short calyx lobes, pubescent outside, tube
2.5–3 mm long; standard blade glabrous, suborbicular, 11.0–11.6 × 10.5–11.5 mm, reflexed, emarginate at the apex,
with a yellow-green patch and a 2.5–3.0 mm long claw; wing blade glabrous, oblong, 8.7–9.0 × 3.5 mm, with a 3.0–3.5
mm long claw; keel blade pocketed at side, pubescent on the outer surface at apex, oblong, 8.8–9.0 × 3.8–4.5 mm, with
a 3.0–3.5 mm long claw; stamen 10, monadelphous, 11.5–12.0 mm long, with two small openings (fenestrae) at base;
floral disk indistinct; ovary appressed hairy, with 5–6 ovules; style inflexed, glabrous; stigma minutely capitate. Pods
compressed at first and becoming inflated with time, slightly to deeply constricted between seeds, 4.2–15.5 × 2.2–5.1
× 2 cm, pubescent, with a distinct beak at apex, without warts outside, usually indehiscent; sutures thickened; fruiting
pedicels 0.4–0.6 cm long, ca. 1.5 mm wide. Seeds 1–4, large, dark brown, reniform, 3–3.7 × 2.6–2.8 cm.

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Phenology and habitat:—Flowering from March to April, and fruiting from April to next March. The flowers
and leaves develop together. It grows in evergreen forests at elevations of 600–1500 m.
Distribution:—China and Thailand. The species is first reported here for the flora of Thailand (Fig. 5).
Additional specimens examined.
CHINA. Yunnan: Cangyuan, 16 Jun. 1974, Y. H. Li 12254 (HITBC), 680–700 m, 19 Jun. 1974, Y. H. Li 12352
(HITBC, KUN, SYS); Jiangcheng, 22 Jun. 2011, S. S. Zhou 10147 (HITBC), 846 m, 19 June 2020, D. P. Ye 919
(HITBC); Jinghong, 755 m, 21 Nov. 2006, T. Zhang et al. SCSB-B-000273 (KUN), 1300 m, 1987, G. D. Tao 14594
(HITBC, IBSC), 750 m, 9 Aug. 1977, G. D. Tao 16715 (HITBC), 1000 m, 21 Apr. 1957, Sino-USSR Exped. 8056
(IBSC); Jinggu, 950 m, 6 Aug. 2001, H. Wang 4886 (HITBC, IBSC); Lancang, 1400 m, 21 Oct. 1989, G. D. Tao &
X. W. Li 39719 (HITBC); Menghai, 1200 m, 29 Oct. 2012, J. W. Li 2683 (HITBC), 1179 m, 31 Mar. 2021, Z. Q. Song
2021019 (IBSC); Mengla, 900 m, 31 Oct. 1983, Exped. 23833 (HITBC), 963 m, 20 Aug. 2020, Z. Q. Song 202039
(IBSC), cultivation, 19 Aug. 2020, Z. Q. Song 202033 (IBSC); Pu’er, 800–1300 m, 20 September 1955, P. I. Mao
6140 (KUN), 900 m, 14 Aug. 1977, G. D. Tao 17384 (HITBC); Shuangjiang, 1050 m, 13 Nov. 1981, Anonymous 2035
(SWFC); Zhenkang, 1500 m, Mar. 1936, C. W. Wang 72150 (A, KUN, LBG). THAILAND. Chiang Mai: Chiang Dao
District, 1400 m, 27 Sep. 1994, W. Nanakorn et al. 1846 (HITBC, IBSC).
2. Derris taiwaniana (Hayata) Z.Q. Song, comb. nov.
≡ Pongamia taiwaniana Hayata (1913: 79) ≡ Millettia taiwaniana (Hayata) Hayata (1920: 22) ≡ Whitfordiodendron taiwanianum (Hayata)
Ohwi (1936: 660). Type:—CHINA. Taiwan, Taibei, Sankakuyu, 1902, K. Nagai s.n. (syntype, not seen); CHINA. Taiwan, 13 March,
S. Yokoyama 27 (syntype, K001414935!).
= Millettia pachycarpa Bentham (1852: 250), non Derris pachycarpa Merrill (1922: 312). Type:—INDIA. Upper Assam. Jenkins
s.n. (Lectotype: K001415586!, designated here; isolectotypes: K001415584!, K001415585!, L0019105!, NY!, P02141837!,
P02141838!).
= Millettia dunnii Merrill (1918: 139). Type:—CHINA. Guangdong, Boluo, Loh Fau [Luofu] Mountain, in thickets near So Liu Koon, 200
m, 13 August 1917, E. D. Merrill 10861 (holotype: PNH, may be destroyed, photo in A!; isotypes: CAS0007371!, NY00016340!,
US00003978!).
= Millettia fooningensis Hu (1955: 360). Type:—CHINA. Yunnan, Foo-ning [Funing County], 550 m, 11 April 1940, Chi-Wu Wang
88378 (holotype: PE00022404!; isotypes: KUN0754399!, KUN0754400!, KUN0754601!, KUN0754602!, KUN0754603!,
KUN0755303!).
Liana large, climbing. Stems cylindric, dark brown, densely scattered with white to brown lenticels, pithy inside; young
shoots yellow tomentose, glabrescent when mature. Stipules ovate-triangular, ca. 3.5 × 3.0 mm. Leaves imparipinnate,
13–17-foliolate (usually 13), juvenile at anthesis, reddish when young; rachis 21.8–38.8 cm long, including petiole 8.6–
14.0 cm long; leaflet blades oblanceolate to oblong, 5.8–20.3 × 2.3–6.1 cm (ratio usually 2–4), papery to subleathery,
glabrous above, appressed pubescent to densely hairy beneath, acute to acuminate at apex, cuneate to rounded at base;
lateral veins 10–16 on each side of midvein, obvious, looped near margin; petiolules ca. 5 mm long; stipels absent.
Pseudoracemes 10.9–39.8 cm long, usually inserted in the leafless axil of leaf on the lower part of branchlets of current
year; rachis appressed hairy; brachyblasts distinct, wart-like or knob-like, up to 7–11 mm long on the lower part, with
3–5 flowers on the top of each brachyblast; bracts inserted at base of brachyblasts and pedicels; bracteoles 2, inserted at
the base of calyx. Flowers pink, pale purplish white to white, ca. 2.1–2.6 cm long; pedicels 0.7–1.2 cm long, appressed
tawny hairy; calyx campanulate, dark red, 5-lobed, pubescent outside, with short calyx lobes, the lower lobe longest,
ca. 2 mm long, 4 mm wide, the tube ca. 6 mm long; standard blade glabrous, suborbicular, ca. 2.3 × 2.1 cm, reflexed,
emarginate at the apex, with a yellow-green patch and a 3.5–4.5 mm long claw; wing blade pubescent on the outer
surface at apex, oblong, ca. 1.8 × 0.7 cm, with a 7 mm long claw; keel blade pocketed at side, pubescent on the outer
surface at apex, oblong, 1.6 × 0.7 cm, with a 7 mm long claw; stamen 10, monadelphous, 2.2 cm long, with two small
openings (fenestrae) at base; floral disk indistinct; ovary appressed hairy, 2.2 cm long, with 6–8 ovules; style inflexed,
glabrous; stigma minutely capitate. Pods inflated, slightly to deeply constricted between seeds, 6.0–14.6 × 3.2–4.1 ×
3 cm, glabrous when mature, with a distinct beak at apex, densely covered with warts, usually indehiscent; fruiting
pedicels 0.9 cm long. Seeds 1–5, large, dark brown, reniform, 2.1–3.3 × 1.8–2.4 cm.
Phenology and habitat:—Flowering from March to June, and fruiting from April to next March. The flowers and
leaves develop together. It grows in evergreen forests at elevations of 0–2500 m.
Distribution:—Bhutan, China, India, Laos, Myanmar, Nepal, Thailand and Vietnam (Fig. 5).
Notes:—Transfer of Millettia pachycarpa Bentham (1852) to Derris Lour. would lead to a later homonym because

MILLETTIA PACHYCARPA AND M. ENTADOIDES TO DERRIS Phytotaxa 531 (3) © 2022 Magnolia Press • 239
of the existence of Derris pachycarpa Merrill (1922: 312). Pongamia taiwaniana Hayata (1913) is the next earliest
legitimate name for this species. Therefore, according to Art. 11.4 of the ICN (Turland et al. 2018), we made the new
combination Derris taiwaniana (Hayata) Z.Q. Song based on Pongamia taiwaniana Hayata. This new combination is
not threatened by the nomen nudum “Derris taiwaniana Matsum.”, which is merely cited as a synonym of Pongamia
taiwaniana, and thus, an invalid name (Art. 36.1, Turland et al. 2018).
Additional specimens examined.
BHUTAN. Punakha District: 1350 m, 11 Oct. 1984, I. W. J. Sinclair & D. G. Long 5598 (E, K); Tashigang: 1360 m,
18 Jun. 1979, A. J. C. Grierson & D. G. Long 2039 (E, K), 1050 m, 28 Jun. 1979, A. J. C. Grierson & D. G. Long 2350
(E), 3000 ft, 17 Aug. 1915, R. E. Cooper & A. K. Bulley 4504 (BM, E). CHINA. Chongqing: Banan, 19 Nov. 1940,
C. Pei 7722 (NAS); Dazu, 100 m, 12 Jul. 1978, Anonymous 440 (SM); Fengdu, 590 m, 12 May 1964, J. A. Wang & Y.
X. Wang 59 (CDBI); Hechuan, 11 May 1959, 1-Section 1586 (CDBI); Nanchuan, 500 m, 30 Oct. 1957, G. F. Li 65000
(IBSC, KUN, NAS), 800 m, 12 Oct. 1995, S. R. Yi 15680 (MO), 750 m, 21 Aug. 1990, Z. Y. Liu 12864 (IMC), 200 m,
20 Aug. 1996, Z. Y. Liu 2029201 (IMC); Rongchang, 27 May 1978, Anonymous 30 (SM); Tongliang, 500 m, 27 Aug.
1978, Anonymous 549 (SM); Wulong, 24 Sep. 1978, Anonymous 1224 (SM); Fujian: Dehua, 1250 m, 5 May 1930, P.
C. Tsoong 193 (AU, IBSC, PE); Fuzhou, 2 Jun. 1951, T. J. Liu 112 (FJSI), Huaan, 10 Jun. 1959, S. M. Huang 5118
(IBSC), 710 m, 15 Apr. 1987, W. D. Han 20425 (NF), Huai Pin Hsiang, 16 Apr. 1937, H. Migo s.n. (NAS), 11 Jul. 1937,
H. Migo s.n. (NAS); Longyan, 260 m, X. F. Zeng ZXF13580 (CZH); Nanan, 850 m, 31 May 1965, Fujian Exped. 2164
(NAS), Nanjing, 400 m, 26 Feb. 1989, H. B. Chen 2252 (FJSI), 450 m, 10 Apr. 1991, H. B. Chen 2553 (MO), 17 May
1942, J. He 1938 (FJSI, PE); Shanghang, 340 m, 5 Sep. 1983, Meihuashan Exped. 9 (FJSI); Xiamen, 27 Oct. 2019, Z.
W. Mao 363 (AU); Yanping, 280 m, 24 May 1980, H. Y. Zou 477 (NF), 280 m, 21 Oct. 1980, H. Y. Zou 858 (NF), 14
May 1905, S. T. Dunn 2564 (IBSC); Yongchun, 702 m, 24 Oct. 2018, X. F. Zeng ZXF41394 (CZH); Yongtai, 203 m, 7
Nov. 2012, Q. Tian et al. TQ02522 (CSH), 230 m, 30 May 1931, Y. Lin 316 (AU, IBSC, PE); Guangdong: Boluo, 125
m, 13 Aug. 1917, C. O. Levine 1371 (A), Feb. 27 1930, H. T. Ho 60155 (IBK, NY, IBSC), 30 Jul. 1930, N. K. Chun
41461 (IBK, IBSC, SYS), 26 Feb. 1930, S. P. Ko 50112 (NY, IBSC); Chaoan, 690 m, 6 Dec. 2009, X. F. Zeng 8719
(CZH), 510 m, 3 May 2009, X. F. Zeng ZXF6573 (CZH); Conghua, 24 Nov. 1958, Sino-Germany Exped. 1158 (IBSC);
Dabu, 500 m, 2 Nov. 1996, B. H. Chen et al. 83 (IBSC), 350 m, 11 May 1984, Dabu Exped. 489 (IBSC); Dongguan,
2500 ft, 16–17 Mar. 1932, S. Y. Lau 20102 (A, NY, IBSC, SYS); Gaozhou, 6 May 1929, Y. Tsiang 2163 (IBK, IBSC,
SYS); Guangzhou, 27 May 1924, To & Tsang 12170 (A, BM, E, MO, NAS, P, US), 16 May 1935, Y. Tsiang 392 (IBK);
Heping, 298 m, 1 Jul. 2002, C. M. Tam Y06622 (JJF, SZG); Heyuan, 14 Apr. 1930, C. L. Tso 21552 (IBSC); Huidong,
24 May 1983, B. Y. Chen et al. 461 (IBSC); Jiaoling, 650 m, 25 May 1957, L. Teng 4898 (IBSC, NAS, PE); Lechang,
450 m, 4 Aug. 1986, B. Y. Chen et al. 2626 (IBK, IBSC), 8 May 1929, C. L. Tso 20268 (NY, IBSC), 8 May 1929, C. L.
Tso 20271 (IBSC), 17 May 1934, S. P. Ko 54519 (IBK, IBSC, NAS), 300 m, 24 May 1934, S. P. Kuo 80676 (IBSC),
23 Aug. 1950, T. S. Chu 104 (IBSC); Liannan, 18 May 1951, T. S. Chu 60986 (IBSC); Lianzhou, 27 May 1951, T. S.
Chu 730 (IBSC); Longchuan, 24 Jul. 1990, B. H. Chen 992 (IBSC); Longmen, 250 m, 17 May 1986, Nanling Exped.
2155 (IBSC); Maoming, 280 m, 30 Apr. 1957, Zhanjiang Exped. 4087 (IBSC); Nanxiong, 21 Aug. 1985, Z. Y. Li 739
(MO); Qujiang, 14 Jun. 1985, Z. Y. Li 624 (MO); Raoping, 16 Apr. 1931, N. K. Chun 42677 (IBK, IBSC); Ruyuan, 20
Aug. 1935, C. S. Chung 10867 (IBSC), 25 Oct. 1938, S. K. Lau 29096 (IBK, IBSC), 21 Jul. 1941, S. K. Lau 29618
(IBSC); Shenzhen, 9 Dec. 1999, F. W. Xing & Y. X. Zhang 12126 (IBSC), 400 m, 17 Oct. 1992, J. F. Chen 2275 (SZG),
100–200 m, 25 Apr. 2005, S. Z. Zhang et al. 587 (PE, SZG), 100–200 m, 10 May 2005, S. Z. Zhang et al. 1291 (PE,
SZG); Shixing, 150 m, 11 Aug. 1958, L. Teng 7166 (IBSC, NAS), 27 Jul. 1985, X. B. Ye 35010 (MO); Wengyuan, 22
Sep. 1933, S. K. Lau 2382 (A, SYS), 23 Nov. 1939, S. K. Lau 25293 (IBK, IBSC); Xinfeng, 1–19 Jun. 1938, Y. W. Taam
859 (P, IBSC); Xinxing, 3 Oct. 1958, Y. S. Lau 2434 (IBSC); Xinyi, 18 Jul. 1931, C. Wang 30979 (IBK, IBSC, SYS),
25 Apr. 1932, C. Wang 32211 (IBK), 29 Nov. 1951, T. S. Chu 1137 (IBSC); Yangchun, 4 Nov. 1935, C. Wang 38641
(IBK, IBSC), 300 m, 6 Sep. 1990, Yunkai Exped. 46 (IBSC); Yangshan, 29 Jul. 1936, L. Teng 206 (IBSC), 400 m, 17
Sep. 1985, Nanling Exped. 1463 (IBSC), Jul. –Sep. 1932, T. M. Tsui 762 (A, MO, NY, IBSC, NAS, PE, SYS); Yingde,
18 Oct. 1931, Anonymous 61400 (IBSC), 11 May 1931, H. Y. Liang 60953 (IBK, IBSC, PE), 14 Jan. 1929, Y. K. Wang
510 (A, IBSC, K, MO, P, PE), 9 Jun. 1985, Z. Y. Li 576 (MO); Yunfou, 10–22 Feb. 1928, Y. Tsiang 1892 (NAS); Yunfu,
26 Mar. 1955, C. Wang & L. Teng 501 (IBSC), 19 Nov. 1934, L. Teng 10116 (IBSC); Zhaoqing, 6 Jun. 1975, G. L. Shi
11457 (IBSC), 9 Aug. 1977, G. L. Shi 13089 (IBSC); Guangxi: Baise, 20 Apr. 1936, Kwangsi Prov. Mus. 11188
(IBSC), 2000 ft, 12 Sep. 1928, R. C. Ching 7359 (A, NY, IBSC, NAS); Bama, 26 Apr. 1957, Y. K. Li P01014 (IBK);
Cangwu, 200 m, 21 Jul. 1956, S. H. Chun 9954 (IBSC), 200 m, 21 Jul. 1956, S. H. Chun 9956 (IBK, KUN, LBG);
Chongzuo, 251 m, 27 Mar. 2015, Y. T. Peng et al. 451402150327042LY (GXMG); Dahua, 355 m, 23 Apr. 2018, D. X.
Nong et al. 451029180423017LY (GXMG); Daxin, 29 Jul. 2016, Y. T. Peng et al. 451424160729013LY (GXMG), 26

SONG & PAN
240 • Phytotaxa 531 (3) © 2022 Magnolia Press
Jul. 1958, Z. X. Zhang et al. 3842 (IBK); Debao, 613 m, 26 Jun. 2015, Debao Exped. 451024160626122LY (GXMG,
IBK); Donglan, 200 m, 18 Jan. 1958, C. C. Chang 11364 (IBK, IBSC); Du’an, 21 Apr. 1978, Du’an Exped. 4-10-0232
(GXMI), 5 Jul. 1957, Y. K. Li P01613 (IBK); Fengshan, 933 m, 2 Apr. 2013, B. Y. Huang et al. 451223130402046LY
(GXMG), 696 m, 25 Aug. 2013, D. X. Nong et al. 451223130825037LY (GXMG), 750 m, 26 Oct. 2012, H. Z. Lv et al.
451223121026048LY (GXMG); Fusui, 24 Apr. 1957, S. H. Chun 12047 (IBK, IBSC, KUN); Guanyang, 209 m, 26
Nov. 2015, Guanyang Exped. 450327151126006LY (GXMG, IBK); Guilin, 28 Oct. 1950, C. S. Chung 808606 (IBK,
IBSC, PE); Hechi, 11 Jun. 1928, Anonymous 92059 (IBK); Hengxian, 30 Apr. 1957, Z. Z. Chen 50352 (IBK); Hexian,
200 m, 20 Oct. 1989, Daguishan Exped. 81028 (GXMI); Huanjiang, 500 m, 23 Oct. 1991, Dian-Qian-Gui Exped.
70158 (IBK), 236 m, 20 Jul. 2013, Huangjiang Exped. 451226130720003LY (GXMG, IBK); Jingxi, 26 Jul. 1977, Y.
Lin 3-54276 (GXMI); Jinxiu, 24 Feb. 1954, Anonymous 24 (IBK), 18 Apr. 1982, Dayaoshan Exped. 14381 (IBSC),
310 m, 19 Apr. 1982, Dayaoshan Exped. 14411 (MO, IBSC), 500 m, 22 Nov. 1981, Dayaoshan Exped. 811983 (GXMI),
9 May 1929, S. S. Sin 8222 (IBSC), 28 Jun. 1934, S. S. Sin 23356 (IBK, IBSC), 11 Dec. 1958, Y. C. Chen 1163 (IBK);
Laibin, 3 Aug. 1977, Y. A. Shi 574-92 (GXMI); Leye, 5 May 1960, Lingle Exped. 33022 (IBK), 1250 m, 14 May 1960,
Lingle Exped. 33115 (IBK), 23 Jun. 1960, N. K. Liang 10173 (GXMI), 19 Jun. 1960, N. K. Liang 11115 (GXMI), 882
m, 10 Sep. 2013, X. Y. Huang et al. 451028130910030LY (GXMG); Lingchuan, 615 m, 17 May 2013, Lingchuan
Exped. 450323130517016LY (GXMG, IBK); Lingui, 12 Jan. 1953, L. H. Chun 93247 (IBK, MO, IBSC, PE), 189 m,
11 Aug. 2015, Lingui Exped. 450322150811023LY (GXMG), 12 May 1964, X. Z. Zheng 249 (GXMI); Lingyun, 18 Jul.
1933, A. N. Steward & H. C. Cheo 711 (P), 21 Dec. 1958, C. T. Ting & J. Z. Wang 1336 (NAS), 17 Apr. 1957, C. Wang
43053 (IBSC), 8 Dec. 1984, Guangxi Exped. 327 (GXMI), 350 m, 10 Sep. 1989, Huanan Exped. 1180 (IBSC), 450 m,
8 Oct. 1989, Huanan Exped. 2338 (IBSC), 617 m, 21 Mar. 2013, Lingyun Exped. 451027130321036 (GXMG, GXMI),
14 Jul. 1937, S. K. Lau 28639 (IBK, IBSC, KUN, PE); Longan, 24 Apr. 1978, X. X. Chen & Y. P. Huang 2-281 (GXMI);
Longlin, 650 m, 24 Jul. 1959, Anonymous 604 (HGAS), 24 Jul. 1959, Anshun Exped. 604 (KUN), 800 m, 5 Nov. 1957,
C. C. Chang 10812 (IBK, IBSC, KUN), 1 May 1957, C. F. Liang & T. L. Wu 32033 (IBK, IBSC), 560 m, 14 Oct. 1957,
Nanzhidi 4593 (IBK, IBSC), 1000 m, 10 Sep. 1987, S. Q. Tang et al. 130 (IBK); Longsheng, 480 m, 13 May 2014,
Longsheng Exped. 450328140513016LY (GXMG), 3 May 1987, X. X. Chen & D. R. Liang 6133 (GXMI); Longzhou,
550 m, 23 Dec. 1957, P. C. Tam 57590 (IBSC); Luocheng, 21 Apr. 1978, Luocheng Exped. 4-1-086 (GXMI), 179 m,
21 Apr. 2013, Luocheng Exped. 451225130421004LY (IBK); Nandan, 2500 m, 22 Jun. 1937, C. Wang 40840 (IBK,
IBSC, PE); Nanning, 25 Mar. 1959, D. Fang 1907 (GXMI); Napo, 1400 m, 9 Dec. 1958, C. C. Chang 13506 (IBK,
IBSC), 19 Oct. 1962, C. C. Lee 1215 (IBK), 22 Dec. 1958, C. T. Li 602312 (IBK, PE), 1250 m, 4 Jan. 1959, C. T. Li
602533 (IBK), 1200 m, 23 Apr. 1981, D. Fang et al. 22420 (GXMI), 10 Oct. 1935, S. P. Ko 55892 (IBSC); Ningming,
12 Oct. 1958, C. C. Chang 12167 (IBK, IBSC, KUN), 9 Jun. 1959, H. Q. Li 40911 (IBK), 15 Nov. 1959, X. F. Deng
10591 (IBK); Pingguo, 13 Oct. 2015, H. Z. Lv et al. 451023151013061LY (GXMG); Pingle, 22 Dec. 1958, S. Q. Zhong
A62834 (KUN, PE); Pingxiang, 377 m, 15 Apr. 2016, Y. Y. Xie et al. 451481160415005LY (GXMG); Qinxian, 330 m,
1 Aug. 1958, C. C. Chen 490 (IBSC); Quanzhou, 344 m, 30 Jan. 2013, Quanzhou Exped. 450324130130027LY
(GXMG, IBK); Rongshui, 700 m, 5 Jun. 1959, Liuzhou Exped. 2346 (IBK); Shanglin, 19 Apr. 1978, Shanglin Exped.
2-432 (GXMI), 20 Apr. 1978, Shanglin Exped. 2-439 (GXMI); Shangsi, 13 Jul. 1959, H. Q. Li 40865 (IBK), 30 m, 7
Jan. 1944, S. H. Chun 4288 (IBSC); Tiandeng, 730 m, 29 Apr. 2015, H. Z. Lv et al. 451425150429044LY (GXMG);
Tiandong, 17 Sep. 1977, S. G. Wang 3-15226 (GXMI), 146 m, 26 Mar. 2018, Tiandong Exped. 451022180326024LY
(GXMG); Tian’e, 23 Aug. 1958, C. T. Li 601326 (IBK, KUN), 17 Jul. 1977, Tiane Exped. 4-6-613 (GXMI); Tianlin,
200 m, 5 Jun. 1958, C. T. Li 600614 (IBK, IBSC, KUN), 16 Jun. 1936, H. Y. Liang 67725 (IBK, IBSC), 600 m, 18 Apr.
1989, Hongshuihe Exped. 89-69 (PE), 320 m, 21 Mar. 2013, Tianlin Exped. 451029130321001 (GXMG), 1072 m, 22
Apr. 2013, Tianlin Exped. 4510291300422040 (GXMG); Tianyang, 21 Nov. 2002, N. Li et al. 11280 (SZG), 676 m, 23
May 2016, Tianyang Exped. 451021160523031LY (GXMG); Tze-an, 1400 ft, 3 Jul. 1928, R. C. Ching 6341 (A, NY,
US, IBSC, NAS, PE); Wuming, 21 Apr. 1978, G. G. Xie 2-106 (GXMI); Xingan, 400 m, 16 May 2016, Xingan Exped.
450325160516005LY (GXMG); Yangshuo, 9 May 1954, C. S. Chung 31292 (IBK), 280 m, 1 Nov. 1963, Z. Z. Chen
53271 (IBK); Yao-shan, S. S. Sin 592 (IBSC); Yongfu, 425 m, 25 Mar. 2013, Yongfu Exped. 450326130325030LY
(GXMG, IBK); Yulin, 26 May 1959, Y. K. Li 404459 (IBK); Zhaoping, 26 Oct. 1958, Y. K. Li 403556 (IBK, IBSC).
Guizhou: Anlong, 1000 m, 12 May 1960, Guizhou Exped. 2393 (NAS, PE), 500 m, 14 May 1960, Guizhou Exped.
2616 (NAS), 1100 m, 9 Jun. 1960, Guizhou Exped. 3185 (HGAS, NAS), 700 m, 16 May 1960, Guizhou Exped. 3449
(HGAS), 1200 m, 15 Jun. 1960, Guizhou Exped. 3534 (HGAS), 1300 m, 20 May 1960, Guizhou Exped. 3646 (HGAS),
900 m, 24 May 1960, Guizhou Exped. 3741 (HGAS), 1100 m, 12 Jun. 1960, Guizhou Exped. 4343 (HGAS), 1700 m,
15 Jun. 1960, Guizhou Exped. 4534 (NAS), 28 Sep. 1985, J. B. Zuo 75-007 (GFS); Bijie, 648 m, 22 Nov. 2016, S. Y.
Peng 522401161122009LY (GZTM); Ceheng, 600 m, 3 May 2005, F. C. Wang 637 (HITBC, PE), 4 Nov. 1930, Y.
Tsiang 9204 (A, NY, IBSC, NAS), 23 Oct. 1980, Z. L. Ding & H. H. Zhang 25 (GFS); Chishui, 500 m, 10 Apr. 1959,

MILLETTIA PACHYCARPA AND M. ENTADOIDES TO DERRIS Phytotaxa 531 (3) © 2022 Magnolia Press • 241
Bijie Exped. 1162 (KUN), 850 m, 25 Sep. 1959, Bijie Exped. 1744 (KUN), 430 m, 27 Jul. 1998, D. X. Wang 2073
(GFS), 28 Mar. 1986, H. H. Zhang 1204 (GFS), 5 Apr. 1986, H. H. Zhang 1322 (GFS), 500 m, 22 Jul. 1995, Z. T. Wang
& Z. Y. Cao 57 (PE); Congjiang, 430 m, 7 May 1982, J. M. Yuan 838 (HGAS); Daozhen, 393 m, 13 May 2016, Pucha
Sect. 520325160513545LY (GZTM), 650 m, 7 Jun. 2003, Z. Y. Liu et al. 2032994 (IMC); Dejiang, 720 m, Guizhou
Uni. Exped. DJ-0135 (GZAC), 493 m, 2 May 2016, M. C. Wang 520381160502097LY (GZTM), 473 m, 21 May 2016,
X. G. Lu 522227160521025LY (GZTM); Dushan, 984 m, M. T. An DS-0561 (GZAC), 330 m, 25 Aug. 1930, Y. Tsiang
6679 (A, IBSC, NAS); Guanling, 24 Nov. 1935, S. W. Teng 1675 (IBSC); Guiding, 1908, J. Cavalerie 3340 (K);
Huangping, 1299 m, 23 Mar. 2017, M. C. Wang 520402170323407LY (GZTM); Jinping, 9 Jun. 1965, J. M. Wang 1196
(GFS); Kaiyang, 610 m, 23 Sep. 1983, M. Z. Yang 1570 (HGAS), 900 m, 24 Sep. 1983, M. Z. Yang 1664 (HGAS);
Libo, 25 Jul. 1959, Libo Exped. 1384 (HGAS, PE), 730 m, 8 May 1981, M. Z. Yang 810274 (HGAS), 350 m, 2 May
1983, X. H. Song 657 (NF), 650 m, 22 May 1983, X. H. Song 857 (NF), 780 m, 1 Oct. 1983, X. H. Song 1111 (NF), 350
m, 2 May 1983, Xiang-Hou Song et al. 657 (K, MO), 650 m, 22 May 1983, Xiang-Hou Song et al. 857 (K), 545 m, 14
May 1982, Y. K. Li 9427 (HGAS, PE); Liping, 350 m, 16 Jun. 1981, D. F. Huang 766 (HGAS, NAS); Luodian, 200 m,
27 Mar. 1960, Guizhou Exped. 44 (HGAS), 180 m, 29 Mar. 1960, Guizhou Exped. 375 (HGAS), 621 m, 2 Nov. 2015,
Luodian Exped. 522728151102005LY (GZTM), 1026 m, 23 Oct. 2015, M. C. Wang GYZ20151023017 (GYBG), 600
m, 6 Apr. 1959, S. Guizhou Exped. 257 (HGAS, KUN), 200 m, 8 Apr. 1959, S. Guizhou Exped. 368 (KUN), 10 Apr.
1959, S. Guizhou Exped. 766 (HGAS); Pingtang, 22 Apr. 1965, Anonymous 31 (HGAS), 700 m, 22 Apr. 1965,
Anonymous 74 (HGAS); Renhuai, 10 Oct. 1929, P. C. Tsoong 146 (PE); Rongjiang, 480 m, 2 Aug. 1965, Z. P. Jian et
al. 51357 (HGAS, KUN); Sandu, 630 m, 29 Aug. 2004, J. Y. Ping 31 (HITBC, PE), 6 Jan. 1960, S. Guizhou Exped. 21
(HGAS), 500–303 m, 27 Oct. 1980, Y. K. Li et al. 8541 (HGAS, IBSC, KUN); Suiyang, 1123 m, 3 Jun. 2015, M. C.
Wang GYZ201506030103 (GYBG); Taijiang, 12 May 2001, M. T. An 210 (GFS); Tongzi, 200 m, 7 Feb. 1931, Y. Tsiang
8054 (A, NY, IBSC, NAS); Wanmo, 28 Jun. 1973, Anonymous 79 (HGAS), 750 m, 21 Jun. 1965, Anonymous 843
(HGAS), 800 m, 11 Jul. 1981, D. J. Liu 129 (HGAS), 400 m, 19 Apr. 1960, Guizhou Exped. 1813 (HGAS), 19 Jun.
1999, M. T. An 991064 (GFS); Weining, 2100 m, 22 Jun. 2014, M. C. Wang YD760622-554 (GYBG); Wudang, 1200
m, 17 Aug. 2014, Wudang Exped. 52011213100403LY (GZTM); Xingren, 14 Aug. 1960, Guizhou Exped. 8188 (HGAS,
PE); Xingyi, 7 Aug. 1959, Anshun Exped. 876 (KUN), 1141 m, 11 Jul. 2015, C. Y. Deng 522301150711684LY (GZTM),
1050 m, 14 Jul. 1960, Guizhou Exped. 6165 (HGAS), 15 Jul. 1960, Guizhou Exped. 6165 (HGAS), 1300 m, 12 Jul.
1960, Guizhou Exped. 6992 (KUN, NAS), 10 Aug. 1960, Guizhou Exped. 7471 (HGAS), 800–1000 m, 23 Jun. 1917,
Handel-Mazzetti 118 (A); Xishui, 850 m, 25 Sep. 1959, Bijie Exped. 1746 (HGAS, KUN); Yanhe, 850 m, 10 Sep. 2003,
J. W. Xiao 5834 (HITBC, PE); Zhenan, 1400 m, 15 Oct. 2015, H. Liu 520324151015016LY (GZTM); Zhenning, 1 Mar.
1993, L. Yang 149 (GNUG); Zhijin, 1103 m, 11 Nov. 2015, B. B. Luo 522425151111010LY (GZTM), 14 Aug. 1975, Y.
L. Tu 338 (GNUG); Ziyun, 1124 m, 19 Jun. 2017, M. C. Wang GYZ201706190008 (GYBG), 1237 m, 08 Jun. 2017, X.
Q. Hou 520425170608351LY (GZTM); Zunyi, 783 m, 17 Mar. 2016, P. F. Zhang 522121160317019LY (GZTM);
Hunan: Baojing, 19 Oct. 2012, D. G. Zhang 1019036 (JJU), 3 Jul. 2009, D. G. Zhang 907030535 (JJU), 500 m, 16
Sep. 1958, L. H. Liou 9819 (KUN, IBSC, PE), 320 m, 14 Nov. 2012, X. J. Su & H. B. Li 433125D00181114000 (JJU);
Dong’an, 660 m, 7 Oct. 2004, J. K. Liu 767 (PE), 500 m, 15 Jul. 2002, X. L. Yu et al. 2121 (CSFI); Guzhang, 334 m, 7
Oct. 2018, C. B. Ma & L. J. Shi HY20180710-0389 (JJU), 253 m, 13 Apr. 2017, D. G. Zhang & D. F. Zhang
LX20170413017-0017 (JJU); Jianghua, 6 Aug. 1958, C. J. Qi 3691 (CSFI); Jiangyong, 800 m, 9 Jul. 1959, P. C. Tam
62239 (IBK), 350 m, 11 Aug. 1997, X. L. Yu 1323 (CSFI), 250 m, 25 Apr. 1984, Y. T. Xiao 40584 (CSFI); Jishou, 18
May 2008, H. S. He 2008051808 (JJU), 193 m, 3 Nov. 2018, S. Xiang JS20180311-0168 (JJU); Longshan, 1 Aug.
2013, Y. Xiao et al. LS-2464 (CSH); Luxi, 15 Aug. 2017, D. G. Zhang ZZ170815852 (JJU); Sangzhi, 200 m, Sep. 1963,
C. J. Qi 30324 (CSFI); Shuangpai, 420 m, 30 Aug. 2002, X. L. Yu & Q. H. Xu 1878 (CSFI); Suining, 440 m, 7 Jul. 2013,
J. J. Zhou & D. Zhou 13597 (CSFI); Tongdao, 250 m, 13 Sep. 1984, K. W. Liu 32815 (CSFI), 29 May 1988, T. C. Chen
580 (IBSC); Yanling, 550 m, Nov. 1988, K. W. Liu 33867 (CSFI); Yizhang, 14 Mar. 1941, P. H. Liang 83066 (IBK,
IBSC), 200 m, 26 Jan. 1942, S. H. Chun 90 (IBSC, PE), 105 m, 12 May 1942, S. H. Chun 1013 (IBK, IBSC), 12 Jul.
1942, S. H. Chun 1871 (IBK, IBSC); Yongshun, 220–270 m, 2 May 1988, Beijing Exped. 778 (PE), 16 Oct. 1990, D.
G. Zhang 4443 (JJU), 24 Jul. 1959, S. C. Lee 204581 (IBK, IBSC); Jiangxi: Dayu, 600 m, 10 Jul. 1965, Anonymous
9450 (KUN); Xunwu, 260 m, 3 Oct. 2016, X. F. Zeng ZXF22192 (CZH), 305 m, 7 Jan. 2017, X. F. Zeng ZXF24228
(CZH); Sichuan: Anyue, 490 m, 24 May 1978, Anonymous 1180 (SM); Changning, 820 m, 20 May 1959, Anonymous
560 (PE); Dechang, 1800 m, 15 May 1959, S. G. Wu 870 (CDBI, KUN); Ebian, 28 Oct. 1938, T. N. Liou 12666 (PE);
Emeishan, 22 Jun. 1931, C. L. Zhou 6289 (HGAS), 1 Sep. 1939, C. W. Yao 4942 (PE), 1 Sep. 1939, C. W. Yao 4949
(NAS), 30 Sep. 1938, Cheo & Tsu 726 (NAS), 600 m, 31 Aug. 1957, G. H. Yang 57036 (IBSC, KUN, NAS, PE), 600
m, 26 May 1995, H. G. Xu 1995434 (MO), 13 Oct. 1952, J. H. Xiong et al. 33133 (IBK, IBSC, NAS, PE), 18 Oct. 1952,
J. H. Xiong et al. 33274 (IBK), 900 m, 9 Oct. 1957, K. C. Liu 3132 (CDBI), 6 Oct. 1939, T. N. Liou & C. Wang 1335

SONG & PAN
242 • Phytotaxa 531 (3) © 2022 Magnolia Press
(PE, WUK), 7 Jun. 1941, W. P. Fang 17478 (PE), 480 m, Nov. 1941, W. P. Fang 18038 (IBSC, JXU), 9 Aug. 1935, X.
B. Zhang 319 (NAS), 750 m, Nov. 1928, Y. Chen 7570 (NAS, NF), 27 Jul. 1935, Y. Y. Ho 5496 (NAS), 30 Nov. 1946,
W. K. Hu 9167 (E, US), 22 Jan. 1929, W. P. Fang 6393 (K, SYS); Gulin, 800 m, 2 Jul. 1976, Anonymous 279 (SM);
Huili, 1 Apr. 1914, C. Schneider 627 (K), 1550 m, 1 Apr. 1914, Handel-Mazzetti 1064 (E); Junlian, 500 m, 22 Jul.
1977, Anonymous 1075 (SM); Leibo, 29 May 1959, Anonymous 443 (PE); Leshan, 200 m, 3 Sep. 1959, C. T. Kuan
6123 (PE), 400 m, 19 Nov. 1934, T. T. Yü 4368 (K), 400 m, 19 Nov. 1934, T. T. Yü 4368 (PE), 15 Sep. 1981, Z. X. Li et
al. 7 (CDBI); Mabian, 8 May 1931, F. T. Wang 22823 (PE), 700 m, 2 Apr. 2000, H. G. Xu 240043 (MO), 600 m, 25
May 1978, Mabian Sect. 179 (SM); Miyi, 7 Jul. 1958, S. Y. Chen et al. 10649 (NAS, SM); Miyi, 1500 m, 22 Aug. 1958,
Z. He et al. 11370 (NAS, SM); Muchuan, 800 m, 15 Aug. 1999, Y. F. Xu 180050 (MO); Nanchong, 500 m, 29 Apr.
1958, Nanchong Exped. 133 (CDBI, KUN, PE); Pingshan, 800 m, 1 Jun. 1959, Anonymous 743 (PE), 770 m, 6 Jun.
1934, T. T. Yü 1919 (K), 600 m, 6 Jun. 1934, T. T. Yü 2939 (PE), 600 m, 6 Jun. 1934, T. T. Yü 2939 (K), 13 May 1930,
W. P. Fang 170 (A, E, K, NY, US, IBSC, PE); Weiyuan, 400 m, 18 May 1978, Weiyuan Sect. 77 (SM); Xinjin, 600 m,
22 Jul. 1938, T. P. Wang 8291 (WUK, PE); Xuyong, 475 m, 15 May 1959, Luzhou Exped. 797 (CDBI), 22 Aug. 2013,
W. B. Ju & X. N. Deng HGX12364 (CDBI), 470 m, 15 Mar. 2013, X. F. Gao et al. HGX11278 (CDBI); Ya’an, E. H.
Wilson 4608 (A, K), 1000 m, 2 Sep. 1999, Y. F. Xu 190020 (MO); Yingjing, 23 Nov. 1951, W. G. Hu & Z. He 12093
(PE); Taiwan: Pingdong, 1 Jan. 1917, Matuda Eizi 1056 (TAI); Taibei, 11 Oct. 1918, E. H. Wilson 10783 (A), 300–500
m, 28 Dec. 1984, S. F. Huang 2303 (TAI); Tainan, 700–800 m, 11 Dec. 1992, H. F. Yen 7197 (MO); Taizhong, 500–600
m, 21 Sep. 1990, T. C. Huang 14784 (TAI); Taoyuan, 520 m, 16 Sep. 1996, C. M. Wang & H. M. Lin 2191 (MO),
500–800 m, 9 Apr. 1990, J. M. Hu & K. C. Yang 748 (A, MO), 2030 m, 28 Mar. 1997, M. F. Loa 193 (PE); Xinzhu,
1000 m, 17 Jan. 1992, C. C. Liao & C. C. Wang 146 (A, MO), 565 m, 13 Jan. 2003, C. M. Wang 6470 (MO, PE), 30
Dec. 1932, S. Sasaki s.n. (MO); 13 Mar., S. Yokoyama 27 (K); Xizang: Chayu, 1500 m, 13 Jul. 1980, Z. C. Ni et al. 602
(XZ); Motuo, 650 m, 14 Sep. 1974, Anonymous 2530 (PE), 1100 m, 12 Apr. 1983, B. S. Li & S. Z. Cheng 4037 (PE),
800 m, 12 Oct. 1992, H. Sun et al. ETM-0389 (KUN), 1400 m, 12 Nov. 1992, H. Sun et al. ETM-1252 (KUN), 1000
m, 25 Feb. 1993, H. Sun et al. ETM-3965 (KUN), 1400 m, 25 Feb. 1993, H. Sun et al. ETM-4019 (KUN), 1400 m, 16
Oct. 2015, J. Hu et al. HJ05282 (NAS), 1000 m, 6 Aug. 1974, Kezi Exped. 1656 (KUN, PE), 800 m, 11 Aug. 1974,
Qinghai-Xizang Exped. 74-4197 (PE); Yunnan: Baoshan, Apr. 1914, G. Forrest 12353 (BM, E, K, IBSC); 7000–8000
ft, Apr. 1921, G. Forrest 19371 (A, E, K); Bijiang, 1200 m, 15 Jul. 1978, Bijiang Exped. 1286 (KUN); Cangning, 1330
m, 28 Sep. 1988, Anonymous 1039 (KUN); Cangyuan, 1700 m, 28 Mar. 2005, S. S. Zhou 2527 (HITBC); Daguan, 700
m, 21 Jun. 1973, B. Y. Sun et al. 738 (KUN, PE); Eshan, 1500 m, 7 Jul. 1989, Yuxi Exped. 89-295 (KUN); Fengqing,
1900 m, 4 Oct. 1958, J. Chen 128 (KUN), 1200 m, 21 Nov. 1977, Q. Lin 7730 (KUN), 1800 m, 13 May 1938, T. T. Yü
15859 (KUN); Fugong, 1300 m, 2 Jun. 1978, Bijiang Exped. 342 (KUN); Fugong, 5 Nov. 2011, S. S. Zhou 11037
(HITBC); Funing, 1000 m, 1 Nov. 1964, Q. A. Wu 9696 (KUN), 14 Jun. 1964, S. Z. Wang 971 (KUN); Gongshan, 1600
m, 30 May 1950, Anonymous 10044 (KUN), 1400 m, 11 Jul. 1979, Q. Lin & X. F. Deng 790826 (KUN), 1300 m, 2 Oct.
1965, 65-Wenshan Exped. 16 (KUN); Guangnan, 1100 m, 14 Jun. 1964, S. Z. Wang 1063 (KUN); Hekou, 23 Dec.
1974, Hekou Exped. 93 (KUN, PE); Huaning, 1507 m, 24 Dec. 2012, C. Liu et al. 12CS4767 (KUN); Jingdong, 2300
m, 4 Nov. 1956, P. Y. Chiu 53427 (KUN), 1200 m, 8 Nov. 1963, Z. H. Yang et al. 101483 (KUN), 1000 m, Aug. 1936,
C. W. Wang 76363 (A, IBSC, NAS, PE), 900 m, 18 Aug. 1977, G. D. Tao et al. 17659 (KUN, HITBC), 9 Apr. 2011, S.
S. Zhou 8684 (HITBC); Lincang, 1300 m, 23 Aug. 1957, S. S. Sin 270 (KUN, IBSC, PE); Lushui, 840 m, 2 Apr. 1957,
W. X. Liu 133 (KUN, PE), 150 m, 7 Arp. 1957, W. X. Liu 165 (KUN, PE); Menghai, 1000 m, 4 Apr. 1996, B. G. Li &
L. Sun 960268 (HITBC), 1300 m, 9 Mar. 1957, Sino-USSR Exped. 7151 (KUN); Mengla, 300 m, 12 Apr. 1955, K. M.
Feng 20248 (KUN, IBSC, PE), 620 m, 19 Sep. 1961, Y. H. Li 3571 (KUN, HITBC), 900–1000 m, 2 Nov. 1974, Z. H.
Yang et al. 12131 (HITBC); Mengzi, 6000 ft, A. Henry 10521 (A, K, MO, NY, US); Mojiang, 1717 m, 2 Apr. 2021, Z.
Q. Song 2021023 (IBSC), 1697 m, 2 Apr. 2021, Z. Q. Song 2021027 (IBSC); Nanjian, 1390 m, 13 Jun. 2012, Z. F. Xu
et al. 3732 (KUN); N’mai kha-Salwin divide, 26°N, 7000–8000 ft, May 1919, G. Forrest 17858 (E, K); Pingbian, 760
m, 21 Apr. 1954, P. I. Mao 3943 (WUK, PE); Pu’er, 4500 ft, 16 Apr., A. Henry 13000 (A, E, K, MO, NY), 4000 ft, A.
Henry 13530 (A, K, K, MO, NY, US), 1500 m, 5 Apr. 2000, H. Wang 3890 (HITBC), 1000 m, 2 Oct. 1955, Kunming
Station 6384 (KUN); Shizong, 12 Nov. 1984, H. Sun et al. 53 (KUN), 800 m, 15 Nov. 1984, H. Sun et al. 137 (KUN),
970 m, 17 Aug. 1985, S. C. Ho 85326 (IBSC), 800 m, 3 Jul. 1987, Xiangke Exped. 870381 (KUN); Shuangbai, 1750
m, 19 Apr. 1957, W. Q. Yin 704 (KUN, WUK); Shuangjiang, 1600 m, 19 Sep. 1957, S. S. Sin 1045 (KUN, IBSC);
Shweli-Salwin divide, 25°20’ N., 7000–8000 ft, Aug. 1912, G. Forrest 9030 (E, PE), 8000–9000 ft, Jun. 1919, G.
Forrest 17891 (E); Tengchong, 5000 ft, May 1912, G. Forrest 7576 (E, K, PE), 5000 ft, Jul. 1912, G. Forrest 8570
(BM, E, K, PE); Jun. 1913, G. Forrest 12031 (E), 5000 ft, Jun. 1913, G. Forrest 12081 (BM, K), 1580 m, 5 Oct. 1977,
P. H. Yu 20227 (HITBC); Weishan, Weishan Exped. 5329271085 (IMDY); Wenshan, 1350 m, 5 May 1962, K. M. Feng
22450 (KUN, WUK); Yangbi, 29 Mar. 1933, H. D. McLaren C176 (A, BM, E), 7 Nov. 1941, T. N. Liou 22611 (PE), 6

MILLETTIA PACHYCARPA AND M. ENTADOIDES TO DERRIS Phytotaxa 531 (3) © 2022 Magnolia Press • 243
Mar. 1988, Y. H. Li et al. 541 (KUN); Yanjin, 560 m, 1 Sep. 1964, N. E. Yunnan Exped. 968 (KUN); Yiliang, 450 m,
24 Sep. 1972, N. E. Yunnan Exped. 974 (KUN, PE); Yongde, 840 m, 6 Jul. 2005, E. D. Liu et al. 1090 (KUN);
Yongping, 1500 m, 20 Oct. 1985, Q. Su 309 (KUN); Yuanjiang, 1380 m, 6 Nov. 1964, Y. H. Li 5894 (KUN, HITBC),
1500 m, 4 Jul. 2012, Yuanjiang Exped. 5304280853 (IMDY); Zhenxiong, 550 m, 8 Sep. 1987, Xiangke Exped. 870596
(KUN); Zhejiang: Taishun, 250 m, 21 Apr. 1990, Li et al. 428 (ZM). INDIA. Assam: Cachar, R. L. Keenan s.n. (K);
Grwhatly, 29 Mar. 1886, C. B. Clarke 43242D (BM); Jeypore, 1898, Prain Collector s.n. (A); Meghalaya: Khasia
Hills, 2000 ft, 2 Apr. 1886, C. B. Clarke 43280A (BM); Griffith Kew Dist. 1767 (K, P); Nagaland: Naga Hills, 5000
ft, Dec. 1907, A. E. Meebold 7123 (K, K), 7000 ft, 4 Sep. 1950, W. N. Koelz 26069 (L); Kala Naga Hills, 4000 ft, May
1882, G. Watt 7257 (E, K, L); Sikkim:1–3000 ft, J. D. Hooker s.n. (A, BM, K, L, P); West Bengal: Choowbultee,
Darjeeling, 12 Jun. 1870, C. B. Clarke 12274 (BM); Darjeeling, 2500 ft, May 1875, J. S. Gamble 2102A (K); Mangbar,
3000 ft, Apr. 1904, H. H. Haines 2029 (E, K); Selimpahar, 3000 ft, 30 Jun. 1876, J. S. Gamble 765A (K); Swoke, 11
Arp. 1910, Ribu & Rhomoo 4081 (L); Tista, 1000 ft, 10 Apr. 1923, G. H. Cave s.n. (A, E); W. Duars, 11 Feb. 1879, J.
S. Gamble 6669B (K). LAOS. Phongsali: Phou Den Dinh Phinh Ha, 27 Apr. 1936, M. Poilane 25904 (P); Xieng
Khouang: Phou Khe, 1500 m, 3 Apr. 1952, J. E. Vidal 1479 (P). MYANMAR. Chin State: Natma Taung National
Park, 887 m, 12 Mar. 2014, P. Srisanga et al. 97919 (E); Kachin State: Myitkyina District, 3100 ft, 11 Apr. 1927,
Maung Mya 5365 (K); Myitkyina District, 4000 ft, 3 Apr. 1919, R. Farrer 844 (E); Nansau, 500 ft, Apr. 1911, M. Kyaw
41 (E); Mandalay Region: Maymyo, 3700 ft, 18 Apr. 1917, C. G. Rogers 881 (E), 3500 ft, 13 Jul. 1908, J. H. Lace
3289 (E, K); J. H. Lace 5821 (E, K), 3500 ft, 22 Apr. 1913, J. H. Lace 6162 (A, E, E, K); Tanintharyi: Tenasserim,
4000 ft, 14 Apr. 1877, G. Gallatly 725 (BM). NEPAL. Morang District: 26.48N, 87.55E, 1000 ft, 20 Apr. 1971, J. D.
A. Stainton 6805 (BM). THAILAND. Chiang Mai: Chiang Dao District, 26 Sep. 1971, J. E. Vidal 5189 (P); Doi Phra
Kao, 1470 m, 1 Sep. 1931, H. B. G. Garrett 714 (K, L); Mae Rim District, 1300 m, 16 Jun. 2009, J. F. Maxwell 09-163
(L); Mae Wang District, 1275 m, 20 Mar. 2003, J. F. Maxwell 03-46 (A, L); Me Chem, 900 m, 12 May 1921, A. F. G.
Kerr 5423 (BM, K, TCD); Me Kong, 1190 m, 2 Apr. 1925, Winit 1318 (K); Muang District, 1610 m, 15 Sep. 1989, J.
F. Maxwell 89-1087 (A, E, L, MO), 1600 m, 22 Mar. 1991, J. F. Maxwell 91-289 (A, E, L, P); Mueang Chiang Mai
District, 5200 ft, 9 Apr. 1911, A. F. G. Kerr 1770 (BM, K, TCD); Sameung, 1350 m, 20 Jul. 1997, J. F. Maxwell 97-787
(A, L, KUN); Chiang Rai: Mae Sai District, 550 m, 20 Aug. 2004, I. C. Nielsen et al. 1856 (L, P); Phan District, 575
m, 10 Sep. 1997, J. F. Maxwell 97-1007 (A, L); Lampang: Mueang Pan District, 550 m, 4 Dec. 1995, J. F. Maxwell
95-1303 (L); Mae Hong Son: Pai District, 1450 m, 6 Apr. 1990, J. F. Maxwell 90-396 (A, E, L, MO); Phayao:
Mueang Phayao District, 650 m, 7 May 1997, J. F. Maxwell 97-483 (A, L); Ranong: Mueang Ranong District, 10 m,
22 Mar. 1993, Sino-Thai Botanical Exped. 1335 (KUN). VIETNAM. Cao Bang: Cho Ra, Phan Ke Loc P7052 (HN);
Lai Chau: Phong Tho, 600 m, 6 Apr. 1936, M. Poilane 25533 (P); Lang Son: Huu Lung District, 100 m, D. K. Harder
& P. K. Loc 4089 (MO, PE); Ninh Binh: Cuc Phuong, 12 Jul. 1998, Chin Cuong Ha 98798 (CPNP).
Acknowledgements
We thank the curators and staff of herbaria A, AU, BM, CDBI, CPNP, CSFI, CSH, CZH, E, FJSI, GFS, GNUG, GXMG,
GXMI, GYBG, GZAC, GZTM, HGAS, HITBC, HN, IBK, IBSC, IMC, IMDY, JJF, JJU, JXU, K, KUN, L, LBG, MO,
NAS, NF, NY, P, PE, SM, SYS, SZG, TAI, TCD, US, WUK, XZ, and ZM for hosting our visits or providing images of
related specimens online. We thank the Herbarium of the Arnold Arboretum of Harvard University for permission to
use the image in this paper. We also thank Prof. Richard T. Corlett for the improvement of the manuscript. This study
was supported by the National Natural Science Foundation of China (grant no. 31600165).
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Appendix 1.
Sampled taxa and the GenBank accessions (trnK/matK, ITS/5.8S, trnL-F IGS, psbA-trnH IGS) for sequence data used
in the study. Missing sequence data are indicated by an n-dash (─). * indicates the sample was newly sequenced in
this study.
Abrus precatorius, JN407123, JN407458, ─, ─; Aganope balansae, JX506601, JX506433, JX506489, JX506544;
Aganope gabonica, JX506605, JX506438, ─, JX506548; Aganope heptaphylla, JX506600, JX506432, JX506488,
JX506543; Aganope impressa, JX506604, JX506436, JX506492, JX506547; Aganope stuhlmannii, JX506603,
JX506435, JX506491, JX506546; Aganope thyrsiflora, JX506602, JX506434, JX506490, JX506545; Antheroporum
glaucum, LC080895, ─, ─, ─; Antheroporum pierrei, KX713080, ─, ─, ─; Apios americana, KF279516, KF272967, ─,
─; Austrosteenisia blackii, AF142707, AF467020, ─, ─; Canavalia africana, KT751466, KT751420, ─, ─; Canavalia
albiflora, KT751467, KT751421, ─, ─; Craibia brevicaudata, JX517315, ─, ─, ─; Craibia lujai, KX713087, ─, ─,
─; Craibia zimmermannii, JX518072, ─, ─, ─; Brachypterum cumingii, JX506618, JX506447, JX506505, JX506561;
Brachypterum eriocarpum, JX506625, JX506454, JX506512, JX506568; Brachypterum involutum, JX506622,
JX506451, JX506509, JX506565; Brachypterum koolgibberah, JX506624, JX506453, JX506511, JX506567;
Brachypterum microphyllum, JX506619, JX506448, JX506506, JX506562; Brachypterum philippinense, JX506627,
JX506455, ─, ─; Brachypterum pseudinvolutum, JX506623, JX506452, JX506510, JX506566; Brachypterum robustum,
JX506617, JX506446, JX506504, JX506560; Brachypterum scandens, JX506621, JX506450, JX506508, JX506564;
Brachypterum submontanum, JX506626, ─, JX506513, JX506569; Brachypterum thorelii, JX506620, JX506449,
JX506507, JX506563; Dahlstedtia muehlbergiana, JX506615, AF467059, JX506502, JX506558; Dahlstedtia pinnata,
MN076275, MN076216, ─, ─; Deguelia negrensis, JX506607, JX506441, ─, ─; Deguelia sp., JX506608, JX506440,
JX506495, JX506551; Derris alborubra, JX506638, JX506466, JX506524, JX506580; Derris amoena 1, JX506628,
JX506456, JX506514, JX506570; Derris amoena 2, JX506629, JX506457, JX506515, JX506571; Derris amoena
3, JX506630, JX506458, JX506516, JX506572; Derris cuneifolia, JX506649, JX506478, JX506535, JX506591;
Derris elegans, JX506641, JX506469, JX506527, JX506583; Derris elliptica C, JX506647, JX506475, JX506533,
JX506589; Derris elliptica K1, JX506648, JX506477, JX506534, JX506590; Derris elliptica K2, ─, JX506476, ─, ─;
Derris elliptica ST, JX506646, JX506474, JX506532, JX506588; Derris ferruginea, JX506633, JX506461, JX506519,
JX506575; Derris fordii H77*, OK345322, MZ871610, OK345326, OK345318; Derris glabra, JX506635, JX506463,
JX506521, JX506577; Derris laotica, JX506645, JX506473, JX506531, JX506587; Derris laxiflora, AF142715,
AF467046, ─, ─; Derris lianoidnes, JX506653, JX506482, JX506539, JX506595; Derris luzoniensis, JX506654,
JX506483, JX506540, JX506596; Derris marginata, JX506643, JX506471, JX506529, JX506585; Derris montana,
JX506650, JX506479, JX506536, JX506592; Derris monticola, JX506637, JX506465, JX506523, JX506579; Derris
oblongifolia, JX506652, JX506481, JX506538, JX506594; Derris piscatoria, JX506651, JX506480, JX506537,
JX506593; Derris pseudomarginata, JX506639, JX506467, JX506525, JX506581; Derris pubipetala, JX506634,
JX506462, JX506520, JX506576; Derris reticulata, JX506632, JX506460, JX506518, JX506574; Derris rubrocalyx,
JX506644, JX506472, JX506530, JX506586; Derris solorioides, JX506640, JX506468, JX506526, JX506582; Derris
spanogheana, JX506636, JX506464, JX506522, JX506578; Derris tonkinensis, JX506631, JX506459, JX506517,
JX506573; Derris trifoliata, JX506642, JX506470, JX506528, JX506584; Fordia cauliflora, HM049511, GQ434352,
─, GU396708; Fordia nivea, MG816795, ─, ─, ─; Fordia splendidissima 1, , AF142718, AF467048, ─, ─; Fordia
splendidissima 2, MN076277, MN076218, ─, ─; Fordia stipularis, MG816797, ─, ─, ─; Hesperothamnus ehrenbergii,
KX713100, KJ411645, ─, ─; Hesperothamnus pentaphyllus, KX652168, KJ411646, ─, ─; Ibatiria furfuracea,
MN076274, MN076215, ─, ─; Indigofera amblyantha, MH658881, MH711821, ─, ─; Indigofera potaninii,
MN722147, MN722001, ─, ─; Indigofera sokotrana, GU951671, EU729559, ─, ─; Kunstleria ridleyi, JX506598,
─, JX506486, ─; Leptoderris brachyptera, JX506611, JX506444, JX506498, JX506554; Leptoderris fasciculata,
JX506609, JX506442, JX506496, JX506552; Leptoderris hypargyrea, JX506610, JX506443, JX506497, JX506553;
Lonchocarpus lanceolatus, AF142717, AF467057, ─, ─; Lonchocarpus leucanthus, MN076276, MN076217, ─,
─; Lonchocarpus santarosanus, JX506613, AF467063, JX506500, JX506556; Lonchocarpus sericeus, JX506612,
JX506485, JX506499, JX506555; Lonchocarpus subglaucescens, JX506614, AF467066, JX506501, JX506557;
Millettia brandisiana, MN076305, MN076219, ─, ─; Millettia caerulea, KY241795, ─, ─, ─; Millettia diptera, ─,
MH156226, ─, ─; Millettia drastica, MN370281, ─, ─, ─; Millettia duchesnei, MN370203, ─, ─, ─; Millettia dura,
MN966639, ─, ─, ─; Millettia entadoides H110*, OK345323, MZ871611, OK345327, OK345319; Millettia extensa 1,
KY241797, ─, ─, ─; Millettia extensa 2, LC080901, ─, ─, ─; Millettia grandis, AF142724, AF467474, ─, ─; Millettia
ichthyochtona, ─, AF467475, ─, ─; Millettia kangensis, KY241798, ─, ─, ─; Millettia lasiantha, ─, AF467476, ─,
─; Millettia laurentii, MZ274110, ─, MZ274110, MZ274110; Millettia leptobotrya, AF142725, AF467477, ─, ─;

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Millettia leucantha, LC080899, ─, ─, ─; Millettia leucantha var. buteoides, KY241800, ─, ─, ─; Millettia leucantha
var. latifolia, KY241799, ─, ─, ─; Millettia macrostachya, KY241801, ─, ─, ─; Millettia makondensis, KF147413,
─, ─, ─; Millettia mossambicensis, JX517618, ─, ─, ─; Millettia pachycarpa, KY241803, MH156227, ─, ─; Millettia
pachycarpa H113*, OK345325, MZ871612, OK345329, OK345321; Millettia pachycarpa H85*, OK345324,
MZ871613, OK345328, OK345320; Millettia pallens, MN370279, ─, ─, ─; Millettia peguensis 1, KY241806, ─,
─, ─; Millettia peguensis 2, KY241805, ─, ─, ─; Millettia pinnata, JX506616, JX506445, JX506503, JX506559;
Millettia pulchra 1, KP093474, KP092735, ─, ─; Millettia pulchra 2, KP093475, KP092736, ─, ─; Millettia rhodantha,
MN370285, ─, ─, ─; Millettia richardiana, ─, AF467483, ─, ─; Millettia richardiana, AF142713, AF467483, ─, ─;
Millettia stuhlmannii 1, JX517411, ─, ─, ─; Millettia stuhlmannii 2, MZ274111, ─, MZ274111, MZ274111; Millettia
tecta, KY241802, ─, ─, ─; Millettia thonningii, AF142723, AF467481, ─, ─; Millettia usaramensis, JX905956, ─,
─, ─; Millettia xylocarpa 1, LC080903, ─, ─, ─; Millettia xylocarpa 2, KY241807, ─, ─, ─; Millettia zechiana,
MN370280, ─, ─, ─; Mucuna macrocarpa, LC494404, LC494605, ─, ─; Mundulea sericea, MN076310, MN076248,
─, ─; Ostryocarpus riparius, JX506599, JX506431, JX506487, JX506542; Philenoptera eriocalyx, AF142720,
AF467487, ─, ─; Philenoptera laxiflora, AF142721, AF467488, ─, ─; Philenoptera laxiflora, ─, ─, JX506494,
JX506550; Philenoptera violacea 1, MN076314, MN076214, ─, ─; Philenoptera violacea 2, JX506606, JX506439,
JX506493, JX506549; Piscidia carthagenensis, JQ587806, KJ411709, ─, ─; Piscidia mollis 1, ─, KJ411710, ─, ─;
Piscidia mollis 2, ─, AF467489, ─, ─; Piscidia piscipula, AF142710, AF467490, ─, ─; Platysepalum chevalieri,
KY118195, ─, ─, ─; Platysepalum hirsutum, ─, EU729482, ─, ─; Platysepalum violaceum, KX713115, ─, ─, ─;
Pongamiopsis amygdalina 1, MN076308, MN076246, ─, ─; Pongamiopsis amygdalina 2, AF142711, AF467494,
─, ─; Tephrosia astragaloides, MN076312, MN076249, ─, ─; Tephrosia benthamii, MN076311, MN076250, ─, ─;
Tephrosia heckmanniana, AF142712, AF467497, ─, ─; Tephrosia luzoniensis, MH767999, MH768294, ─, ─.