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Millettia weizhii (Fabaceae: Millettieae), a new species with four‐winged pods from a limestone area in Yunnan, China

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Millettia weizhii sp. nov. (Fabaceae), a tree species bearing pods with 4 wings is described and illustrated from a limestone area in Yunnan, China. Fruiting specimens of the new species were previously identified as Millettia tetraptera Kurz, a species recorded from Myanmar and India (West Bengal), but examination of specimens and data from our field survey show that the new species is easily distinguished by having longer and stipitate pods, much shorter pseudoracemes with 2 flowers at each node, white and longer corolla, triangular calyx lobes and more and narrower leaflets. The new species should be placed in the Merrillii/Xylocarpa‐group of Millettia sect. Fragiliflorae Dunn based on morphological characters.
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NORDIC JOURNAL OF
BOTANY
Nordic Journal of Botany
1
––––––––––––––––––––––––––––––––––––––––
© 2018 e Authors. Nordic Journal of Botany © 2018 Nordic Society Oikos
Subject Editor: Cheng Du
Editor-in-Chief: Torbjörn Tyler
Accepted 12 December 2018
2019: e01964
do i: 10.1111/njb .0196 4
doi: 10.1111/njb.01964 00 1–6
Millettia weizhii sp. nov. (Fabaceae), a tree species bearing pods with 4 wings is described
and illustrated from a limestone area in Yunnan, China. Fruiting specimens of the new
species were previously identied as Millettia tetraptera Kurz, a species recorded from
Myanmar and India (West Bengal), but examination of specimens and data from our
eld survey show that the new species is easily distinguished by having longer and
stipitate pods, much shorter pseudoracemes with 2 owers at each node, white and
longer corolla, triangular calyx lobes and more and narrower leaets. e new species
should be placed in the Merrillii/Xylocarpa-group of Millettia sect. Fragiliorae Dunn
based on morphological characters.
Keywords: China, Leguminosae, limestone area, tree species
Introduction
Millettia Wight & Arn. is a large genus in the tribe Millettieae (Fabaceae), and its
taxonomy has been considered as a globally problematic (Geesink 1984). Dunn
(1912) conducted a comprehensive revision of the genus and accepted 138 species
in 16 sections. Since then, however, many new species have been described and the
circumscription of the genus has been changed repeatedly. e current status of the
genus is presented in the ‘Legumes of the World’ (Schrire 2005): four sections of
Millettia sensu Dunn have been excluded, viz. sect. Eurybotryae Dunn, sect. Austro-
millettia Dunn, sect. Bracteatae Dunn and sect. Albiorae Dunn (Geesink 1984, Schot
1994, Hu and Chang 2003), while two genera (Pongamia Vent and Neodunnia R. Vig.)
and one section of Lonchocarpus Kunth (sect. Caudaria Dunn) have been transferred to
Millettia (Polhill 1971, Geesink 1981, Labat and Du Puy 1995).
As presently circumscribed, the genus Millettia comprises about 150 (Schrire 2005)
or 216 species (Hesperothamnus Brandegee is excluded, Hu and Chang 2003), which
are distributed in Africa and Asia. e number of Asian Millettia species may be esti-
mated to about 80, including at least 50 species of trees, based on Lock and Heald
(1994), Adema (2000), Lôc and Vidal (2001), Kumar and Sane (2003), Lock and Ford
(2004) and Zhu (2015). In China, after removal of Millettia sapindifolia T. C. Chen
Millettia weizhii (Fabaceae: Millettieae), a new species with
four-winged pods from a limestone area in Yunnan, China
Zhu-QiuSong, QianXia, KaiZhang, BoPan, Jian-WuLi, Dong-XianXu, ShuaiLiao and Shi-JinLi
Z.-Q. Song (http://orcid.org/0000-0002-7763-2431), Q. Xia, K. Zhang and S.-J. Li (lisj@scib.ac.cn), South China Botanical Garden, Chinese Academy
of Sciences, Guangzhou, P. R. China. ZQS, QX and KZ also at: Univ. of Chinese Academy of Sciences, Beijing, P. R. China. – B. Pan and J.-W. Li,
Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, P. R. China. – D.-X. Xu, Guangdong Academy of Forestry,
Guangzhou, P. R. China. – S. Liao, School of Life Sciences, East China Normal Univ., Shanghai, P. R. China.
Research
2
(Songetal. 2017), the genus includes 14 species, 12 of which
are recorded from the Yunnan Province (Zhu 2015).
During examination of herbarium specimens of Chinese
Millettia, we found that some specimens from Yunnan have
pods with 4 wings, diering from other Chinese members of
the genus. Two specimens collected in 1962, Y. H. Li 4100
(HITBC, KUN), were identied by Wei in 1978 as ‘Millettia
tetraptera Kurz’ or ‘Millettia cf. tetraptera Kurz’ (Fig. 1A), and
are thus likely the vouchers for M. tetraptera as reported from
Yunnan by Wei (1985, 1994). Moreover, the line-drawings
provided in Wei (1985, Fig. 5; 1994, pl. 37, Fig. 6) are simi-
lar to the two specimens. is taxon was also recorded from
Yunnan in the subsequent literature (Sun 2006, Zhu etal.
2007, Wei and Pedley 2010, Zhu 2015).
However, after being compared with the protologue of M.
tetraptera, these fruiting materials from Yunnan appear quite
dierent, because they have longer (2327 vs 15 cm) and
stipitate (versus estipitate) pods, and more (usually 1113
vs 57) and narrower (1.53.0 vs 38 cm) leaets. Our eld
survey in Yunnan further indicated that this plant is also dis-
tinct in its type of inorescence, color and length of corolla
and shape of calyx lobes (Table 1). We thus conclude that
this distinct plant from Yunnan has been misunderstood by
previous authors.
Globally, there are four species of Millettia described as
having pods with 4 wings, viz. M. tetraptera, M. glaucescens
Kurz, M. pterocarpa Dunn and M. prainii Dunn (Kurz 1873,
1877, Dunn 1912). e last species has later been united
with M. glaucescens (Grierson and Long 1987, Sanjappa
1992, Pressetal. 2000, Kumar and Sane 2003, Borah 2014).
All these species occur in the tropical regions of Asia (Adema
2000, Kumar and Sane 2003), but morphological compari-
sons (Table 1) revealed that the materials from Yunnan can
be easily distinguished from these species by having more
numerous leaets, much shorter inorescences, white and
longer corolla, triangular calyx lobes, the stipitate ovary and
longer and stipitate pods, and thus represent a new species as
described and illustrated below.
Millettia weizhii Z. Q. Song sp. nov. (Fig. 13)
A species similar to M. tetraptera in the hairiness of leaets
and pods with 4 wings, but easily distinguished by the longer
(2327 vs 15 cm) and stipitate (vs estipitate) pods, much
Figure1. Millettia weizhii sp. nov. (A) Y. H. Li 4100 (HITBC, barcode 0013680), which was identied by Wei in 1978 as ‘Millettia cf.
tetraptera Kurz’, and (B) J. W. Li 2986 (holotype: HITBC, herbarium no. 150677).
3
shorter (35 vs 1520 cm) pseudoracemes with 2 (vs 25)
owers at each node, white (vs pale blue) and longer (2.5
vs 0.91.1 cm) corolla, triangular (vs very short and broad)
calyx lobes, and more (usually 1113 vs 57) and narrower
(1.53.0 vs 38 cm) leaets.
Type: China. Yunnan Province, Mengla County, Mengyuan,
Daoban, 995 m a.s.l., 8 Apr 2013 (fr.), J. W. Li 2986
(holotype: HITBC, herbarium no. 150677, Fig. 1B; isotype:
KUN, barcode 1251989).
Previously referred to as: Millettia tetraptera auct. non Kurz
by Wei (1985, p. 275, Fig. 5; 1994, p. 139, pl. 37), Sun
(2006, p. 413), Zhuetal. (2007, p. 471), Wei and Pedley
(2010, p. 279) and Zhu (2015, p. 226, pl. 4).
Etymology
e new species is named in honor of Prof. Zhi Wei, a
Chinese botanist who made an important contribution to
our knowledge of the family Fabaceae (especially Millettia)
in China.
Description
Trees, up to 20 m tall and 25 cm in diameter at breast
height; bark gray, with a few branches at the base; branches
of the current year ferruginous tomentose, glabrescent;
wood density (dry weight per fresh volume) 0.8 g cm–3.
Buds rounded, tomentose; scales 6–8, broadly ovate. Leaves
(9–)11–13(–15)-foliolate; rachis 8–17 cm, including petiole
0.5–3.0 cm; stipules and stipels absent; leaet blades stiy
papery, opposite or rarely subalternate, 410 × 1.53.0 cm,
with both surfaces yellowish tomentose when young, soon
glabrous adaxially except on midvein, persistently tomen-
tose abaxially during owering, acuminate at apex, rounded
or subcordate and sometimes asymmetric at base; most
leaets narrowly oblong; terminal leaet largest, oblanceo-
late or obovate; the lowest leaet smallest and ovate; prin-
cipal lateral veins 810 on each side of midvein, curved
apically near margins, adaxially impressed, abaxially raised.
Pseudoracemes axillary at top of branchlets, very short,
3–5 cm long, bearing about 10 owers, without distinct
brachyblasts; pulvinus ca 1 mm long at the base of the ino-
rescences; rachis ferruginous tomentose, with 2 owers each
node; bracteoles positioned at the middle of the pedicles,
caducous, linear, 810 × 0.61.0 mm, villous. Pedicels
paired, 0.51.0 cm long, with ferruginous hairs. Flowers
ca 2.5 cm. Calyx campanulate, 5-lobed but upper 2 lobes
united near tip, ferruginous tomentose; lower 3 lobes as long
as tube, triangular, 0.6 cm long, 0.5 cm wide at base. Corolla
white, distinctly veined; standard glabrous, reexed, subor-
bicular, 2.02.2 × 1.82.0 cm, emarginate at apex; claw ca
0.4 cm, with a yellow-green patch at base and 2 basal cal-
losities (i.e. inexed auricles); wing oblong, 2.0 × 0.8 cm,
slightly auriculate at base, obtuse at apex, with a ca 0.5 cm
long claw; keel smaller than wing, 1.8 × 0.8 cm, obtuse at
apex, truncate at base, its lamina falcate, joined, with a ca
0.6 cm long claw. Stamens 10, monadelphous; laments
connate into a tube, with 2 fenestrae at base; anthers basi-
xed, oblong, ca 1.7 × 0.8 mm. Disk absent. Ovary linear,
stipitate, 6-ovuled, clearly thickened on both sutures, vil-
lous only on ventral suture side; stipe 0.4 cm; style curved
upward; stigma capitate. Legume glabrous, linear oblong,
2327 × 3.23.8 cm, at, tapering towards base, obliquely
beaked at apex, tardily dehiscent, stipitate; stipe 2.03.8 cm,
glabrous; fruiting pedicel 12 cm, ferruginous tomentose;
both sutures with two 0.61.2 cm wide woody wings per-
pendicular to them; the bres of the endocarp orientated
at an angle of about 45° with the longitudinal axis of the
legume. Seeds 25.
Table 1. Comparison between Millettia weizhii sp. nov. and three other species of the genus that have pods with 4 wings.
Traits/species M. weizhii M. tetraptera M. glaucescens M. pterocarpa
Leaflet number (9–)11–13(–15) 5–7 7–9 7–9
Leaflet apex acuminate rounded, emarginate or
apiculate
acuminate caudate
Leaflet width (cm) 1.53.0 3–8 2.0–3.5 2.5
Leaflet hairiness both surfaces initially tomentose,
finally glabrous above
both surfaces initially
tomentose, finally
glabrous above
both surfaces glabrous both surfaces
glabrous
Stipule absent present present absent
Inflorescence 35 cm, tomentose 1520 cm, tomentose 1620 cm, almost glabrous 813 cm, almost
glabrous
Each node with 2 flowers with 2–5 flowers with 2 flowers
Flower white, ca 2.5 cm pale blue, 0.91.1 cm steel blue, 1 cm steel blue, 1 cm
Calyx lobes triangular, 0.6 × 0.5 cm very short and broad very short and broad unknown
Ovary with a 0.4 cm stipe sessile sessile sessile
Legume (cm) 2327 × 3.23.8 15 × 3 813 × 3 17 × 2.5
Fruiting pedicle (cm) 12 0.7 0.5 unknown
Fruiting stipe (cm) 2.03.8 sessile sessile sessile
Flowering Jan to Feb Mar to June Mar to May unknown
Distribution China (Yunnan) Myanmar and India (West
Bengal)
Myanmar, Nepal, Thailand
and India (Assam, Sikkim,
West Bengal)
Malaysia (Perak)
4
Distribution, habitat and phenology
Millettia weizhii is known from Yunnan, China (Fig. 4). It
grows in a limestone area at 700–1000 m a.s.l. It is observed
in ower from Jan to Feb and in fruit from Mar to Sep.
Lots of leaves (not all) drop before owering and new leaves
unfold during fruiting. is species may be considered as
‘Endangered’ (EN) under the IUCN (2001) categories and
criteria B2ab(iii).
Figure2. Millettia weizhii sp. nov. (A) habit, (B) abaxial surface of leaets, (C) gray bark, (D) germinating ower bud (red arrow), unger-
minated leaf bud (purple arrow) and the broken top of a branch (blue arrow), (E) pseudoraceme with paired owers, showing the stipe (red
arrow) of a ovary and the pulvinus (purple arrow) at the base of the inorescence, (F) pods with 4 wings, (G) a ower with the dierent
parts separated, showing c = calyx with triangular lobes, s = standard with 2 basal callosities (i.e. inexed auricles), w = wings, k = jointed keels,
m = monadelphous stamens, o1 = stipitate ovary and o2 = magnied part of the ovary with 6 ovules (× 2.5), bar = 1 cm.
5
Similar species
Like other species bearing pods with 4 wings, M. weizhii
should be ascribed to M. sect. Fragiliorae Dunn, because
these species share several common features such as being
trees with estipellate leaves, axillary pseudoracemes, standard
with 2 basal callosities (i.e. inexed auricles), monadelphous
stamens, no disk and an Asian distribution (Dunn 1912).
e basal callosities on the standard blade was considered as
the consistent characteristic of the section by Dunn (1912,
p. 136), while the pulvinus at the base of the pseudoraceme
was viewed as the most striking character of the section by
Geesink (1984, p. 104). Both characters are present also in
the new species (Fig. 2G(s), E, respectively). e species can
be further placed in the Merrillii/Xylocarpa-group of M. sect.
Fragiliorae based on the numerous ovules and seeds (Adema
2000). However, in contrast to the other members of M. sect.
Fragiliorae, the new species has stipitate ovary (Fig. 2E),
which is an important character of another Asian section of
Figure3. Line drawing of Millettia weizhii sp. nov. (A) fruiting branch, (B) inorescence, (C) a single ower, (D) and (E) calyx, (F) stan-
dard, (G) wings, (H) jointed keels, (I) monadelphous stamens and (J) ovary. Drawn from J. W. Li 2986 (fr.) and K. Zhang 007 (.), which
collected from the same individual.
Figure4. Distribution of Millettia weizhii sp. nov. (star, made by the
software ArcGIS 10.2).
6
the genus, i.e. sect. Podocarpae Dunn (Dunn 1912, Geesink
1984). However, M. weizhii can be distinguished from the
latter section by other characters (e.g. the standard with basal
auricles, monadelphous stamens and no disk).
In addition, Hu (1955) reported that another Millettia
species with winged fruits, i.e. M. glaucescens, occurs in
China and provided the image of one owering specimen
(J. Cavalerie 4573, K). Although M. glaucescens was sometimes
listed among unresolved names of Chinese Millettia
(Zhuetal. 2007, Zhu 2015), the relevant voucher specimen
has been shown to represent M. pulchra (Benth.) Kurz (Wei
1985, p. 290). Moreover, we found that two duplicates of
J. Cavalerie 4573 from P (P00552267 and P00552268) were
erroneously identied as Indigofera penduloides Y. Y. Fang &
C. Z. Zheng.
Additional specimens examined (paratypes)
China. Yunnan: Mengla, 780 m a.s.l., 15 May 1962, Y. H. Li
4100 (HITBC, KUN), 900 m a.s.l., 2 Jun 1982, Exped.
32690 (HITBC), 1000 m a.s.l., Sep 1992, H. Wang 1223,
1246 (HITBC), 995 m a.s.l., 21 Dec 2017, Z. Q. Song
2017006 (IBSC), 995 m a.s.l., 31 Jan 2018, K. Zhang 007
(IBSC).
Acknowledgements – We would like to thank the curators of the
herbaria of HITBC, IBSC, KUN, K and P for hosting our visits or
providing high-quality images of some important specimens online
for our studies, Mr Yu-Ming Wang for his help in the eld study,
and Ms Yun-Xiao Liu (IBSC) for preparing the line drawing.
Funding – is work was partly supported by the National Natural
Science Foundation of China (grant no. 31600165, 31670193).
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... After Dunn, Millettia was revised at the regional level by various authors (e.g., Gagnepain 1916, De Wildeman 1920, Craib 1928, Hepper 1958, Gillett 1961, Wei 1985, Adema 2000, Lôc and Vidal 2001, Adomou and Van der Maesen 2002, Mollel and Adema 2006, Wei and Pedley 2010, Adema et al. 2016. However, recent studies indicated that Millettia in its current circumscription is still in need of major revision (Mattapha 2017, 2020, Mattapha et al. 2019a, b, Song et al. 2017a, b, 2019, 2021a. ...
... A full description for this species was provided by Song et al. (2019). ...
... It flowers before leaves expand, from January to February, and fruits from March to September. This species was recently evaluated as 'Endangered' (EN) by Song et al. (2019). ...
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Millettia lantsangensis Z.Wei was described within the tribe Millettieae on the basis of one fruiting and two flowering gatherings from Yunnan Province, China. Our critical examination shows that these gatherings are mixed and represent two distinct species from two different tribes. The fruiting specimens have woody branches, terminal infructescences and inflated pods, closely resembling Callerya sphaerosperma (Z. Wei) Z. Wei & Pedley from the same tribe, while the flowering collections have soft and herbaceous branches, axillary pseudoracemes and wart-like brachyblasts, and belong to Cruddasia insignis Prain from the tribe Phaseoleae. Because one of the flowering specimens was indicated as the type of Millettia lantsangensis, this is here reduced to a synonym of Cruddasia insignis. The genus Cruddasia Prain was previously reported only from Myanmar and Thailand, and is therefore a new genus record for the flora of China.
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Millettia and Derris are two taxonomically complicated genera of the tribe Millettieae (Leguminosae), and when only flowering material is available they are not easily distinguished from each other. A critical examination of literature and specimens as well as a field survey show that Millettia sapindifolia T. C. Chen, a Chinese species described on the basis of two flowering collections, is conspecific with Derris yunnanensis Chun & F. C. How. The two are closely similar in various characters, such as the inflorescence type, monadelphous stamens, branches with longitudinal ridges, and size, shape, number and hairiness of leaflets. Therefore, the former is here reduced to a synonym of the latter.
Article
Changes in botanical names of flowering plants are an issue which comes up from time to time. While there are valid scientific reasons for such changes, it also creates some difficulties to the floristic workers in the preparation of a new flora. Further, all the important monumental floras of the world have most of the plants included in their old names, which are now regarded as synonyms. In north east India, "Flora of Assam" is an important flora as it includes result of pioneering floristic work on Angiosperms & Gymnosperms in the region. But, in the study of this flora, the same problems of name changes appear before the new researchers. Therefore, an attempt is made here to prepare an updated account of the new names against their old counterpts of the plants included in the first two volumes of the flora, on the basis of recent standard taxonomic literatures. In this, the unresolved & controversial names are not touched & only the confirmed ones are taken into account. In the process new names of 470 (four hundred & seventy) dicotyledonous plant species included in the concerned flora are found out.
Article
The genus Millettia Wight & Arn. is revised for the Flora Malesiana area. Two new species are described: M. borneensis and M. glabra. Pongamia velutina (C.T. White) Verdc. is transferred to Millettia; as the epitheton velutina is already in use in Millettia, a new name is proposed: M. velvetina Adema. In total 15 species are recognised for area concerned. A key the species is given. An identification list and an index of names are included.
Article
Morphological characters support the description of five new species of Millettia, M. capuronii Du Puy & Labat, M. hitsika Du Puy & Labat, M. nathaliae Du Puy & Labat, M. orientalis Du Puy & Labat, and M. taolanaroensis Du Puy & Labat, and one new species of Pongamiopsis, P. viguieri Du Puy & Labat, from Madagascar. The genus Neodunnia R. Viguier is considered to be a synonym of Millettia, and two new combinations, M. aurea (R. Viguier) Du Puy & Labat and M. richardiana (Baillon) Du Puy & Labat, are made. /// L'étude des caractères morphologiques permet la description de cinq nouvelles espèces de Millettia: M. capuronii Du Puy & Labat, M. hitsika Du Puy & Labat, M. nathaliae Du Puy & Labat, M. orientalis Du Puy & Labat, et M. taolanaroensis Du Puy & Labat, et une nouvelle espèce de Pongamiopsis, P. viguieri Du Puy & Labat, de Madagascar. Le genre Neodunnia R. Viguier est considéré comme étant un synonyme de Millettia et deux nouvelles combinaisons, M. aurea (R. Viguier) Du Puy & Labat and M. richardiana (Baillon) Du Puy & Labat, sont faites.
Article
'There is indeed so much conformity in the structure of their [Lonchocarpus, Derris, Pongamia, Piscidia, Muellera] flowers and in the general plan of their pods, that I should have had no hesitation in uniting them all into one large genus divided artificially into sections, were it not for the inconvenience, already alluded to, of suppressing long-established and universally recognized genera, when they can be marked out by a tolerably positive character, however artificial and isolated that character may be.' Bentham (i86o: 19) The weight of convention which persuaded Bentham a hundred years ago to retain genera of convenience in this group persists. The reappraisal of illconforming elements presents a continuing problem of whether to weaken already tenuous distinctions, segregate further small genera or make piecemeal transfers to redefined concepts. Mendonga & Sousa (1965) have recently re-segregated the monotypic African genus Capassa Klotzsch for the species more familiarly known as Lonchocarpus capassa Rolfe. Capassa is 'lonchocarpoid' in general facies, but has a distinct wing to the adaxial suture of the fruit. The absence of such a wing is technically the one character which distinguishes Lonchocarpus Kunth from Derris Lour. The species was indeed transferred to Derris by Harms (1902, 1915), but looks so unlike any other species of that genus that inclusion there has not otherwise been considered acceptable. In trying to determine the generic position of this species for the 'Flora of Tropical East Africa', I have noticed that Lonchocarpus capassa is not the only species which threatens the tenuous distinction between Derris and Lonchocarpus, that in the sum of its characters it seems to fall naturally into the distinctive Afro-Madagascan section Paniculati Benth. of Lonchocarpus, and that new generic criteria emerge if certain more obviously extraneous elements are extricated from the complex. I propose to transfer Lonchocarpus section Caudaria Dunn back to Millettia Wight & Arn., transfer the Australian species of Lonchocarpus to Kunstleria Prain and restitute Derris section Aganope (Miq.) Benth. to generic level, including in it Ostryoderris Dunn. With these excisions, Lonchocarpus section Paniculati (including Lonchocarpus capassa) may be easily distinguished from Derris by the thyrsoid panicles, the flowers not fascicled or densely crowded on contracted ultimate branches of the inflorescence. The basic problem of delineating genera in this group remains nonetheless, and I attempt only a provisional patching up of
Article
Two new members of Callerya group in Fabaceae, Endosamara racemosa (Roxb.) Geesink and Callerya vasta (Kosterm.) Schot, are identified based on phylogenetic analyses of chloroplast rbcL sequences. These taxa joined with other previously identified taxa in the Callerya group: Afgekia, Callerya, and Wisteria. These genera are resolved as a basal subclade in the Inverted Repeat Lacking Clade (IRLC), which is a large legume group that includes many temperate and herbaceous legumes in the subfamily Papilionoideae, such as Astragalus, Medicago and Pisum, and is not close to other Millettieae. Endosamara is sister to Millettia japonica (Siebold & Zucc.) A. Gray, but only weakly linked with Wisteria and Afgekia.