Reduction of Blinkworthia (Convolvulaceae) based on multilocus phylogenetic reconstruction and resurrection of a species from synonymy revealed by phenetic analyses

Abstract

Blinkworthia was established as a tropical Asian genus in the morning glory family (Convolvulaceae) with a unique combination of habit, bract, corolla and fruit morphology. Recently, however, investigations of the morphology and anatomy have shown that none of these diagnostic characters can be used to delimit Blinkworthia as an independent genus. This study resolves the generic circumscription using phylogenetic analysis of molecular data from four loci: ITS, trnL-trnF, matK and rps16. The results showed that all members of Blinkworthia were nested within Argyreia, concordant with the current morphological concept of Argyreia. All information robustly supported that Blinkworthia should be subsumed under Argyreia prompting us to make the nomenclatural transfers from Blinkworthia to Argyreia and make the new combinations A. convolvuloides and A. lycioides. In addition, phenetic analyses revealed that a species endemic to Burma, A. campanuliflora, should be resurrected from the synonymy of B. convolvuloides. The three species are circumscribed, described, illustrated and their ecology, distribution and
vernacular names are summarised, all documented with specimen citations. An identification key is provided for these three closely related species.

Full text

Reduction of Blinkworthia (Convolvulaceae) based on multilocus
phylogenetic reconstruction and resurrection of a species
from synonymy revealed by phenetic analyses
Pantamith Rattanakrajang
1,2
, Pirada Sumanon
1
, Paweena Traiperm
1
, George Staples
3
&
Timothy Utteridge
2
Summary. Blinkworthia was established as a tropical Asian genus in the morning glory family (Convolvulaceae) with
a unique combination of habit, bract, corolla and fruit morphology. Recently, however, investigations of the
morphology and anatomy have shown that none of these diagnostic characters can be used to delimit Blinkworthia
as an independent genus. This study resolves the generic circumscription using phylogenetic analysis of molecular
data from four loci: ITS, trnL-trnF,matK and rps16. The results showed that all members of Blinkworthia were nested
within Argyreia, concordant with the current morphological concept of Argyreia. All information robustly supported
that Blinkworthia should be subsumed under Argyreia prompting us to make the nomenclatural transfers from
Blinkworthia to Argyreia and make the new combinations A. convolvuloides and A. lycioides. In addition, phenetic
analyses revealed that a species endemic to Burma, A. campanuliflora, should be resurrected from the synonymy of
B. convolvuloides. The three species are circumscribed, described, illustrated and their ecology, distribution and
vernacular names are summarised, all documented with specimen citations. An identification key is provided for
these three closely related species.
Key Words. Argyreia, classification, generic delimitation, new combination, South-East Asia, systematics.
Introduction
Blinkworthia Choisy is a small genus of Convolvulaceae,
first described with the single species, B. lycioides
Choisy. The genus is distributed throughout south-
eastern Asian from Burma (Prain 1894) through
northern and western Thailand (Gagnepain &
Courchet 1915;Staples2010) to southern China
(Fang & Staples 1995). Blinkworthia was first thought
to be distinguished from the closely related genus
Argyreia Lour. by the involucre of bracts and the
corolla shape, the erect shrub habit and 1-seeded
indehiscent fruit (Choisy 1834,1845; Hallier 1893a).
Since the discovery of the second species,
B. convolvuloides Prain (Prain 1894), a woody climber,
the habit proved unsuitable to use as a generic
character. Recently it has been shown that plants can
switch from climber to erect shrub and vice versa
(Rattanakrajang et al.2018). To date Blinkworthia has
only included two species, B. convolvuloides and
B. lycioides, while a third species, B. discostigma Hand.-
Mazz. (Handel-Mazzetti 1931), has been treated as a
synonym of B. convolvuloides (Fang & Huang 1979).
According to the traditional morphological classifi-
cation (Hallier 1893a), Blinkworthia was included in
tribe Argyreieae Hallier f. of the ‘Echinoconiae’(a
name never formally published), with the genera
Argyreia and Rivea Choisy; Hallier defined this tribe
mainly on the basis of an indehiscent fruit. Later,
Stictocardia Hallier f. was added to the tribe Argyreieae
because it was believed to have an indehiscent fruit
(Hallier 1893b). However, it has been demonstrated
that the fruit of Stictocardia has a unique tardy
dehiscence at the final stage (Austin & Sebsebe
1997). The second tribe in the ‘Echinoconiae’
(Fig. 1), the Ipomoeeae Hallier f., has a dehiscent
fruit; it originally comprised the genera Calonyction
Choisy, Ipomoea L., Lepistemon Blume, and Quamoclit
Mill. (Hallier 1893a); later, Hallier added the genus
Astrochlaena Hallier f. (Hallier 1893b), an illegitimate
name that was replaced by Astripomoea A.Meeuse
(Meeuse 1957).
Toward the end of the 20
th
century, morphological
characters in the two tribes were analysed using
cladistic analysis (Austin 1998); the results differed
from Hallier’s(1893a,1893b) traditional classification.
In Austin’s consensus tree (Austin 1998:Fig.1),
derived from 128 morphological characters analysed
for the 55 known genera of Convolvulaceae,
Accepted for publication 18 August 2022.
1
Department of Plant Science, Faculty of Science, Mahidol University, King Rama VI Road, Bangkok, 10400, Thailand. e-mail: paweena.tra@mahidol.edu
2
Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK.
3
Harvard University Herbaria, 22 Divinity Ave, Cambridge, MA, 02138, USA.
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DOI 10.1007/S12225-022-10052-1
ISSN: 0075-5974 (print)
ISSN: 1874-933X (electronic)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2022

Blinkworthia formed a clade with Argyreia,Lepistemon,
Lepistemonopsis Dammer and Rivea; this clade in turn is
sister to a larger clade comprising Astripomoea,Ipomoea,
Paralepistemon Lejoly & Lisowski, Stictocardia and
Turbina Raf.Austin’s analysis was attempting to show
parallel vs convergent evolution in characters across
the entire family according to the unrooted phenetic
analysis and without consideration regarding tribal
classification. The macro- and micro-morphology of
tribe Argyreieae were next re-investigated using cladis-
tic approaches (Wilkin 1999). In Wilkin’s concept,
Argyreia was grouped with Blinkworthia, therefore,
Argyreia is sister to Blinkworthia sharing a 4-loculed
ovary (Fig. 1; Wilkin 1999).
The placement of Blinkworthia in relation to the
other genera within the tribe Argyreieae remained
unclear. None of the molecular phylogenetic studies
included Blinkworthia (Fig. 1). The first attempt to
explore molecular phylogenetic relationships within
the spiny pollen group by Manos et al. (2001) revealed
the evolutionary relationships between Argyreieae and
Ipomoeeae based on the combined information of
ITS and waxy sequences plus morphological features.
Their results clearly showed two major clades, the first
consisted of Argyreia,Ipomoea in small part, Lepistemon,
Rivea,Stictocardia and Turbina, while the other clade
was composed of Astripomoea and the greater part of
Ipomoea. However, these results demonstrated the
ambiguous status between Ipomoeeae and Argyreieae
because some genera, i.e. Argyreia,Ipomoea and
Turbina, were not monophyletic.
More intensive molecular phylogenetic investiga-
tion of Convolvulaceae re-examined the monophyletic
status of this family based on four conservative regions
of chloroplast DNA: rbcL,atpB,psbE-J, and trnL-trnF
(Fig. 1; Stefanovićet al.2002). This result exhibited
two clades, Argyreieae and Ipomoeeae, within the
spiny pollen grade. However, several genera such as
Blinkworthia,Lepistemonopsis, and Rivea were not in-
cluded in the analysis. Their results showed that tribe
Argyreieae is nested within tribe Ipomoeeae
(Stefanovićet al.2002); the relationship between tribe
Ipomoeeae and Argyreia was paraphyletic. These
results based on cpDNA corroborated the conclusions
of prior researchers (Austin 1998; Wilkin 1999; Manos
et al.2001).
Subsequently, Stefanovićet al.(2003) proposed a
new classification system for the Convolvulaceae
informed by molecular phylogenetic studies and
reviewing the previous data (Manos et al.2001; Miller
et al.2002; Stefanovićet al.2002). In this new
classification, all genera in the historical tribe
Argyreieae were incorporated within the expanded
tribe Ipomoeeae s.l. and this arrangement conformed
to the monophyletic concept of Manos et al.(2001).
Therefore, the tribe Ipomoeeae s.l., the entire spiny-
pollen clade, now comprised ten genera: Argyreia,
Astripomoea,Ipomoea,Lepistemon,Lepistemonopsis,
Fig. 1. Blinkworthia placement over time in six different tribal arrangements and the evidence supporting the placement, organised
chronologically from left to right. Bold indicates tribe or clade names. Dotted lines indicate combinations (or splits); no change in
dashed line, and tribal placement changes in solid line. Genera described after the previous system are marked by an asterisk.
Question marks indicate purported placements.
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Paralepistemon,Rivea,Stictocardia,Turbina and
Blinkworthia. However, Stefanovićet al. (2003) did not
explain the phylogenetic relationship of Blinkworthia
within the Ipomoeeae s.l. because it was not included
in their molecular phylogenetic analysis (Fig. 1).
Blinkworthia was assigned to Ipomoeeae s.l. because
the genus “is likely to be associated with /Argyreiinae
based on its indehiscent fruit with leathery pericarp.”
(Stefanovićet al.2003: 801) and this was postulated to
indicate a close evolutionary relationship. In our view,
this is another instance of ‘agglomerative taxonomy’
sensu Simões & Staples (2017) a practice that has long
plagued studies of Convolvulaceae systematics.
Recently, the generic status of Blinkworthia had
been re-evaluated based on morphological observa-
tions in natural habitats and using anatomical
approaches to characterise flower-to-fruit develop-
ment (Rattanakrajang et al. 2018). In effect, this
study was a test of the traditional characters used for
generic delimitation. The results revealed that none
of the traditional morphological characters can be
used to define the genus nor to delimit Blinkworthia
as distinct from the genus Argyreia. As pointed out
in the conclusion of that study (Rattanakrajang et al.
2018) it remains to use molecular evidence to
resolve taxonomic rank and generic delimitation
issues.
Recently, Blinkworthia and Argyreia were subsumed
under a greatly expanded Ipomoea by Muñoz-
Rodríguez et al. (2019). We are not convinced that
such a sweeping mass-reduction is warranted nor are
we compelled to adopt it without further scrutiny:
alternative taxonomic interpretations for the available
data are possible. For example, several monophyletic
clades are present in the phylogenetic trees presented
in Muñoz-Rodríguez et al. (2019) and with further
study each of these clades might warrant recognition
at generic rank. Indeed, a nomenclatural proposal has
already been published by a group of more than 40
researchers who specialise in Convolvulaceae
(Eserman et al. 2020) that lays the foundation for an
alternative interpretation of the extant phylogenetic
results. We elect not to subsume all spiny pollen
genera under a much-expanded Ipomoea at this time
and accordingly maintain the well-established generic
concepts.
Our aims are to (1) resolve the generic delim-
itation for Blinkworthia with a robust molecular
phylogenetic analysis; (2) test species delimitation
based on two phenetic analyses to clarify whether
two or three species can be recognised; and (3)
provide a taxonomic treatment that resolves the
nomenclature for the species. Here, we offer a
comprehensive molecular phylogenetic analysis of
Blinkworthia and its allies using DNA sequence data
derived from a nuclear region, internal transcribed
spacer (ITS), and three non-coding chloroplast
regions, trnL-trnF,matK and rps16. In addition,
morphological characters were re-examined from
herbarium specimens and field surveys of living
plants in their natural habitats and subjected to
rigorous phenetic analysis.
Materials and Methods
Field collecting and taxonomic methods. Eight collecting
trips were made between 2014 and 2018 to Thailand
and Burma generating 14 new collections of
Blinkworthia and Argyreia; voucher specimen prepara-
tion and deposit were described in Rattanakrajang
et al.(2018). Identifications of the new collections
were made by comparison with specimens in BK, BKF,
and QBG and consultation of taxonomic literature
(Choisy 1834; Prain 1894; Fang & Staples 1995; Staples
2010; Staples et al. 2021). Vouchers for the leaves dried
in silica gel used for DNA extraction as well as
specimens used for the phenetic analysis are cited in
Appendix 1&2.
In total, 120 collections of Blinkworthia were studied
for the taxonomic treatment. Specimens were studied
by PR or GS in BCU, BK, BKF, BM, E, G, HITBC, L, K,
KUN, MBK, NY, P, QBG, and RAF. Digital images for
Chinese specimens in the herbaria GXMI, IBK, IBSC,
NAS, and PE were studied via the Chinese Virtual
Herbarium (CVH 2020) and selected type specimens
were examined as digital images in JSTOR Plants and
other virtual herbaria [K, NY, US, and W]. In later
stages of nomenclatural research, digital images for
selected type specimens were received from CAL and
W. All specimens studied for the taxonomic treatment
are cited in the Additional Specimens Examined
grouped under each species. For the conservation
status, we assessed using the distribution data and
occurrence of populations on the GeoCat website
(Bachman et al. 2011), IUCN guidelines (2022), and
the database on protected areas (UNEP-WCMC 2020).
Phylogenetic analyses. We sampled selected species
from all genera included in the historical tribes
Argyreieae sensu Hallier (1893a) and Ipomoeeae
(Appendix 1). New sequences are reported here for
11 accessions representing Blinkworthia and 14 acces-
sions for other spiny-pollen taxa (Appendix 1).
Additionally, 36 taxon samples were downloaded from
GenBank to complement the species analysed here for
the first time (Appendix 1).
Molecular extraction and amplification using leaf
samples of Blinkworthia spp., Argyreia spp., Lepistemon
binectarifer (Wall.) Kuntze and Rivea ornata (Roxb.)
Choisy preserved in silica-gel and herbarium leaf
fragments were performed with DNeasy Plant Mini
Kit (Qiagen Inc., U.S.A.) to extract total genomic DNA
with a slight modification for the herbarium samples.
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The nuclear, ITS (AB101-AB102) (Carine et al.2004);
and chloroplast, trnL-trnF (c and f) (Taberlet et al.
1991), matK (matK390f-matK1326r) (Hollingsworth
et al.2009) and rps16 (rps16x2F2-trnK (UUU)) (Shaw
et al.2007), non-coding loci were amplified as in
Simões et al.(2015), again with a slight adjustment for
herbarium materials. The amplified products were
purified using the FavorPrep
TM
Gel/PCR Purification
Kit (Favorgen Biotech Corp., Taiwan) following the
manufacturer’s procedure. Each purified fragment
was treated with the same primer for automatic DNA
sequencing (Macrogen Inc., Singapore).
Sequences were initially adjusted using Bioedit
v.7.2.6 (Hall 1999), with all regions subsequently
combined and aligned using default parameters with
MAFFT v.7.305 (Katoh & Standley 2013), with aligned
sequences analysed using three optimality criteria:
Maximum parsimony (MP); Maximum likelihood
(ML); and Bayesian inference (BI).
MP was performed on PAUP* v.4.0b10 software
(Swofford 2003) with deactivation of uninformative
characters selected. The remaining characters were set
as equal weight following the number of potentially
parsimony-informative sites. The tree bisection-
reconnection (TBR) method was performed with
MULTREES option in effect (Felsenstein 1985). Inter-
nal branch support was estimated by parsimonious
bootstrap (BS) using 1,000 replications, and obtained
the consistency index (CI) and retention index (RI)
for the tree.
ML was based on the combined datasets of all loci,
with gaps explained as missing data using RAxML
v.8.2.10 (Stamatakis 2006; Miller et al.2010) on XSEDE
(Towns et al.2014). A starting time was set with a
random topology. As recommend by Miller et al.
(2010), the general-time reversible (GTR) nucleotide
substitution model (Tevaré 1986) was selected as
GTRCAT and GTR+Γ, respectively. For a thorough
ML search for the best tree we performed 1,000
bootstrap replicates.
The jModelTest v.2.1.10 program (Darriba et al.
2012) was used to determine the best-fitting substitu-
tion model of sequence evolution employing Akaike
information criterion (AIC) as the main command
(Akaike 1974) and to suggest a proper model to apply
in BI.
For BI analysis of the concatenated dataset we
applied MrBayes v.3.2.6 (Ronquist et al.2012)on
XSEDE (Towns et al.2014). Six substitution types and
four gamma rate categories were run in matrices with
the proper model, GTR+I+Γ. Two simultaneous Mar-
kov Chain Monte Carlo (MCMC) methods were run
for ten million generations with sampling every 250
generations. We discarded the initial 25% of the
sampled trees as burn-in. The BI consensus tree was
visualised as a representative phylogenetic tree with
statistical support from three different methods, using
the Figtree v.1.4.3 program (Rambaut 2016).
Phenetic analyses. A total of 42 accessions, i.e., 12
accessions of Argyreia campanuliflora, 18 accessions of
Blinkworthia convolvuloides, and 12 accessions of
B. lycioides, were used in the analyses (see Appendix 2).
We provided seventeen principal characters (P1 –
17, Appendix 3), excluding corolla features, which
are lacking in most samples. Characters for our
morphological analyses were selected from the tradi-
tional generic delimitation for Blinkworthia (Choisy
1834; Collett & Hemsley 1890;Prain1894). Six
characters were qualitative (P1, P4, P7, P8, P12,
P14); a standard glossary was followed to reduce bias
(Beentje 2010). The remaining 11 characters were
quantitatively measured with ImageJ v.1.50i (Fig. 2;
Schneider et al.2012). The mean value of each
quantitative character was calculated by at least three
to ten replicate measurements of mature organs. All
character states were separated out in discrete
numbers (Appendix 3).
Six qualitative and eleven quantitative characters
were scored for each operational taxonomic unit
(OTU). A total of 42 accessions were analysed by two
phenetic methods: unweighted pair group method
with arithmetic mean (UPGMA) and principal com-
ponent analysis (PCA). The dendrogram was recon-
structed from NTSYSpc v.2.1m (Rohlf 2000).
Computed information was continually calculated by
themoduleofSAHN.TheUPGMAdendrogramwas
visualised with the Graphics module to exhibit the
correlational clustering differences among all indi-
viduals. The PCA was analysed in Minitab v.16 (2010)
and displayed as a score plot. The values of the
eigenvectors obtained from the first and second
components (PC1 & PC2) of the score plot deter-
mined the primary significant characters for further
statistical analysis.
The statistical analysis was generated using the SPSS
v.18 software package (SPSS, U.S.A.). Descriptive
statistics were calculated initially, listing the means
and standard deviations. An analysis of variance
(ANOVA) was tested to verify significant difference
in each quantitative character which was extracted
from the PCA analysis, and a box plot was also
constructed for more comprehension. Due to unequal
sample sizes of all raw data in this study, the pairwise
differences of population means were indicated by
multiple comparisons and homogeneous subsets
using Tukey's honest significant difference (HSD)
test (Tukey 1953; Kramer 1956). A confidence level
was set at 0.05 for ANOVA and Tukey's HSD test.
After the significant quantitative characters were
obtained from this process, these characters were
selected to construct a key to species.
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Results
Phylogenetic analyses. A summary of molecular variation
found in the combined dataset is provided in Table 1.
The concatenated matrix of one nuclear and three
plastid regions for sixty-four accessions (exceptions are
noted in Appendix 1) comprising 2,565 characters was
used in the analyses. Two hundred and ninety-seven
characters (11.58%) were potentially parsimony-infor-
mative. The Akaike information criterion in the
jModelTest found GTR+I+Γfor the final concatenated
datasets.
Visual inspection for the consensus tree based on
the multilocus datasets, ITS, trnL-trnF,matK and rps16,
was generated from Bayesian Inference analysis. The
four loci supported a clade that showed the difference
of relationship between the out-group and the major
Ipomoeeae clade (Clade A). This result showed the
concatenated dataset contributed the best estimate for
representing evolutionary relationships among
Blinkworthia and other related genera in the tribe
Ipomoeeae. The combined phylogenetic analysis of
nrDNA and cpDNA strongly supported the monophyly
Fig. 2. Ascheme of morphological quantitative measurements of three plant parts, a portion of branch; Bleaf blades; Ca solitary
flower.
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of the Ipomoeeae clade containing two major clades,
/Argyreiinae and /Astripomoeinae, identified and
described previously (Stefanovićet al.2002; Stefanović
et al.2003), [Fig. 3; ML bootstrap (hereafter ML) =
100%; Bayesian Posterior Probability (hereafter PP) =
1.00]. Moreover, it also confirms the same major
clades (B, C, D, E, F, G, H) within the broadly defined
Ipomoeeae (Fig. 3).
Blinkworthia and its placement in Ipomoeeae Clade. All
accessions of the three taxa recognised as Blinkworthia
form a monophyletic clade, here designated as the
Blinkworthia alliance clade (ML = 93%; PP = 0.99, clade
I), which is nested within the larger Argyreia alliance
clade (ML = 100%; PP = 1.00, clade H) including
Argyreia and Rivea (Fig. 3). In addition, a monophyletic
clade of A. campanuliflora was sister to the accessions of
B. convolvuloides (ML = <75%; PP = 1.00), with high
support (ML = 87%; PP = 0.99).
Field work & herbarium study. We could distinguish
three morphologically distinct entities (OTUs) in the
field (living plants) and the herbarium (dried speci-
mens): two of them correspond to Blinkworthia lycioides
and B. convolvuloides. The third entity was initially
thought to be a new (undescribed) species. Further
study of type specimen images disclosed that this third
entity already had a name: Argyreia campanuliflora
(Gage) Thoth., which had been reduced to synonymy
under B. convolvuloides by Staples & Traiperm (2017).
However, the results of the phenetic study presented
below clearly show that there are three distinct taxa,
which are recognised here at species rank.
Phenetic analyses. Qualitative and quantitative charac-
ters were observed and measured from herbarium
specimens and these raw data were analysed with the
UPGMA (Fig. 4) and PCA analyses (Fig. 5). Firstly, the
morphological similarity matrices of the UPGMA
algorithm for Argyreia campanuliflora and two
Blinkworthia species were analysed to reconstruct a
dendrogram (Fig. 4). The dendrogram was obtained
with an R-value of 0.81109 and agreed with the
indicated values. This cluster analysis produced three
well-defined groups by the similarity value at 0.35.
Three distinct clusters comprised A. campanuliflora
and members of Blinkworthia,B. lycioides and
B. convolvuloides, respectively (Fig. 4). For clustering
analysis, the monophyletic cluster of A. campanuliflora
is sister group of Blinkworthia spp.
Secondly, the PCA analysis was performed with 42
accessions of Blinkworthia and Argyreia campanuliflora.
The first two components accounted for 60.9% of the
total variance (44.9% and 60.9%, respectively;
Table 2). The high positive variables from PC1
(greater than 0.3) include the petiole length (P3),
leaf blade width (P5), leaf blade length (P6), outer
sepal width (P15) and outer sepal length (P16). The
leaf blade shape (P4) contributed most to the second
component (Table 2). The high negative variable
from PC1 (less than -0.3) was due to the outer bract
shape (P12) and from PC2 the leaf blade base (P8),
pedicel length (P10), ratio of peduncle length to
pedicel length (P11), bract shape (P12), outer bract
length (P13) and ratio of outer sepal length to outer
bract length (P17) (Table 2).
To visualise the affiliation between the three OTUs,
the PCA analysis was plotted as a scatter-plot (Fig. 5).
The first two principal components separated the
OTUs into three discrete groups, comprising
Argyreia campanuliflora,Blinkworthia lycioides and
B. convolvuloides.
From the primary assessment, the quantitative
characters were analysed and revealed nine charac-
ters which were significantly different among all
OTUs in PCA. These traits were secondarily verified
in ANOVA and the pairwise differences of population
means by multiple comparisons and homogeneous
subsets in the homogeneous comparisons of Tukey's
HSD test (Table 3). All nine characters can signifi-
cantly separate at the 0.05 level in two or three groups
of species (Fig. 6). The group of Argyreia
Table 1. Characterisations of DNA sequence markers used.
DNA
marker
Aligned
length
No. of
accession
Variable
character
Informative
character
Tree
length
Consistency
index (C.I.)
Retention
index (R.I.)
Evolutionary
model
Nuclear
ITS 558 60 113 185 982 0.563 0.777 GTR+Γ
Plastid 2007 59 528 112 1814 0.695 0.758 TPM2f+Γ
trnL-trnF 412 57 92 11 248 0.956 0.943 TPM2uf
matK 764 56 66 46 257 0.911 0.948 TPM2uf+I
rps16 1595 56 159 101 1546 0.649 0.738 TIM2+Γ
Combined dataset 2565 61 371 297 2891 0.627 0.744 GTR+I+Γ
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Fig. 3. Cladogram depicting the Bayesian consensus tree based on the concatenated dataset (ITS, trnL-trnF,matK and rps16) for
the genera Argyreia,Astripomoea,Blinkworthia,Ipomoea,Lepistemon,Rivea,Stictocardia and the out-group, Merremia, under the
GTR+I+Γreversible evolution model. Asterisks (*) indicate all branches with maximum likelihood (ML) bootstrap percent support of
100 and Bayesian inference (BI) of posterior probability (PP) of 1. Any branches without a support value had weak support (<75%
bootstrap or posterior probability of <0.75) in ML or BI analyses. Colour of taxon name indicates the historical tribal assignment for
the species according to Hallier (1893a,1893b): red font signifies tribe Argyreieae; blue font signifies Ipomoeeae s.s. Coloured
boxes highlight taxa discussed in this study. Letters on branches or nodes are clades discussed in the text. The tree is rooted on an
outgroup species, M. hirta.
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Fig. 4. Dendrogram from the UPGMA clustering analysis, representing three clusters of Argyreia campanuliflora, Blinkworthia
lycioides and B. convolvuloides, respectively, and based on similarity coefficient.
Fig. 5. Scatter plot analysed by PCA, the first component in X-axis and the second component on Y-axis, displaying group
dispersion of each OTU: Argyreia campanuliflora (Q1 & Q4), Blinkworthia convolvuloides (Q3 & Q4) and B. lycioides (Q2).
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campanuliflora showed relatively larger size in leaf
blade, outer bract and outer sepal than the others
(Fig. 6A –B, F –H).
All three species were examined morphologically
using herbarium specimens and Table 4summarises
the diagnostic characters useful for species recogni-
tion and the geographic distribution reported on the
specimen labels.
Discussion
All generic characters of Blinkworthia have now been
thoroughly re-evaluated by a variety of methods. Previ-
ously, Rattanakrajang et al.(2018) showed that none of
the morphological characters historically used to define
the genus Blinkworthia were unique: all of them were
shared with the larger genus Argyreia. However, the
previous study did not initiate any taxonomic changes
pending results of a molecular phylogenetic analysis for
Blinkworthia. With those results now in hand we can take
afirm decision and resolve the generic delimitation for
Blinkworthia in this study.
Phylogenetic placement of Blinkworthia within tribe
Ipomoeeae. Our molecular phylogenetic reconstruc-
tion did not support the recognition of Blinkworthia as
a genus, because such a genus would be clearly
paraphyletic with respect to exclusion of Argyreia
campanuliflora. Furthermore, our results showed all
accessions of Blinkworthia and A. campanuliflora nested
Table 2. Generated results of the principal component analysis. Variable characters, eigenvalues and percentages of variances for
first two principal components examined from OTUs of Argyreia campanuliflora and Blinkworthia individuals.Asterisks indicate
significant values.
Representative morphological character
Principal component
12
P1 Habit 0.267 -0.132
P2 Internode length 0.277 -0.173
P3 Petiole length 0.312* 0.070
P4 Leaf blade shape 0.058 0.176*
P5 Leaf blade width 0.340* 0.004
P6 Leaf blade length 0.347* -0.011
P7 Leaf blade apex -0.113 -0.036
P8 Leaf blade base 0.102 -0.356*
P9 Peduncle length 0.293 0.076
P10 Pedicel length 0.094 -0.317*
P11 Ratio of peduncle length to pedicel length -0.145 -0.439*
P12 Bract shape -0.258* -0.349*
P13 Outer bract length 0.208 -0.375*
P14 Outer sepal shape 0.201 0.024
P15 Outer sepal width 0.336* 0.041
P16 Outer sepal length 0.334* 0.063
P17 Ratio of outer sepal length to outer bract length 0.048 -0.470*
Eigenvalue 7.6336 2.7237
Cumulative percentage of eigenvalue 44.9 60.9
Table 3. Multiple comparisons and homogenous subsets of Tukey’s HSD test of the significant quantitative characters in whole
samples.
Character
Species
A. campanuliflora B. convolvuloides B. lycioides p-value
Petiole length (cm) 0.82 ± 0.37
B
0.34 ± 0.12
A
0.34 ± 0.11
A
*
Leaf blade width (cm) 3.16 ± 1.09
C
1.18 ± 0.36
B
0.84 ± 0.30
A
*
Leaf blade length (cm) 8.80 ± 2.90
C
3.30 ± 0.89
B
2.44 ± 0.81
A
*
Pedicel length (cm) 0.21 ± 0.12
B
0.27 ± 0.08
B
0.14 ± 0.71
A
*
Ratio of peduncle length to pedicel length 3.06 ± 0.48
A
14.08 ± 6.72
B
4.54 ± 1.36
A
*
Outer bract length (cm) 0.77 ± 0.25
C
0.63 ± 0.16
B
0.31 ± 0.11
A
*
Outer sepal width (cm) 1.04 ± 0.20
B
0.63 ± 0.12
A
0.57 ± 0.98
A
*
Outer sepal length (cm) 1.52 ± 0.30
B
0.83 ± 0.15
A
0.71 ± 0.19
A
*
Ratio outer sepal length to outer bract length 2.03 ± 0.53
A
3.13 ± 1.04
B
1.52 ± 0.26
A
*
*The differences of average values in the same row are significant at the 0.05 level.
A, B, C
For each character, the Tukey’s HSD test grouping indicates population mean that is also different at a 0.05 level
significantly.
KEW BULLETIN
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Fig. 6. Boxplots showing the significant quantitative characters of Argyreia campanuliflora (left), Blinkworthia convolvuloides
(middle) and B. lycioides (right), respectively, from SPSS: Apetiole length; Bleaf blade width; Cleaf blade length; Dpedicel length;
Eratio of peduncle length to pedicel length; Fouter bract length; Gouter sepal width; Houter sepal length; Iratio of outer sepal
length to outer bract length.
Table 4. Summary of morphological character states and distribution ranges.
Character A. campanuliflora A. convolvuloides A. lycioides
Habit climber climber (rarely erect shrub) shrub (rarely twining)
Venation 5–10 obvious secondary veins 4 –6 obvious secondary veins indistinct secondary veins
Bract shape / size broadly elliptic, boat-shaped / very
unequal
spatulate / ± equal circular or elliptic / very
unequal
Bract position &
orientation
imbricate with calyx, appressed to it,
deeply convex
attached below calyx, reflexed from it,
margins in-rolled
imbricate with calyx,
appressed to it
Pedicel length 1–5mm 1–3mm 1–3mm
Outer sepal
indumentum
both sides glabrous ab a xia l l y sp a rsel y hai r y , ad a xial l y
glabrous
both sides glabrous
Outer sepal shape ovate to orbicular elliptic to ovate or obovate elliptic to ovate
Distribution W Burma S China and N Burma Burma and Thailand
KEW BULLETIN
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within the Argyreia alliance clade with strong support.
Although the Argyreia alliance clade as formed is
paraphyletic without including Rivea species, we do
not discuss this situation until all species of Rivea have
been sampled and resolved in further work. However,
our molecular evidence is certainly congruent with the
previous hypothesis of morphological relationships as
proposed by Rattanakrajang et al.(2018).
Resurrection of Argyreia campanuliflora. After we exam-
ined the living plants in the field, we found that
Blinkworthia convolvuloides is quite distinct morphologi-
cally from Argyreia campanuliflora; the two had been
synonymised by Staples & Traiperm (2017). Addition-
ally, our molecular phylogenetic experiment robustly
supports separation of A. campanuliflora from
B. convolvuloides. Our phenetic and statistical results
based on qualitative and quantitative morphology fur-
ther confirmed that all accessions of A. campanuliflora
are obviously separated as a distinct group from the two
Blinkworthia species. Interestingly, we found that the
bract was the most informative character to discriminate
members in this group of three species. Hence, all
available evidence supports that A. campanuliflora should
be resurrected as a species; Table 4summarises all
diagnostic characters and distribution areas for these
three species.
Taxonomic reduction for Blinkworthia. Our total evi-
dence based on morphology and evolutionary phylo-
genetic framework did not identify a single
synapomorphic character to support diagnosability of
Blinkworthia from the genus Argyreia. Based on these
results we here reduce Blinkworthia to synonymy under
Argyreia and provide a taxonomic account for the
three species recognised, making new combinations
for two of them under Argyreia.
Taxonomic Treatment
Argyreia Lour. (Loureiro 1790: 134), here expanded.
Type species: Argyreia obtusifolia Lour.
Blinkworthia Choisy (Choisy 1834: 430), synon. nov.
Type species: Blinkworthia lycioides Choisy.
Approximately 143 species (including the three
species transferred here) and 5 varieties in tropical
to subtropical Asia (except in New Guinea) and
Madagascar (Traiperm & Staples 2014;Stapleset al.
2015; Traiperm & Staples 2016; Shalini et al.2017;
Staples & Traiperm 2017; Chitchak et al.2018;
Traiperm et al.2019; Shalini et al.2020; Traiperm
& Suddee 2020).
Key to the species
1. Bracts ± equal, spatulate, attached below calyx, reflexed from it, margins in-rolled; outer sepals elliptic to ovate
or obovate, abaxially sparsely hairy, adaxially glabrous . . . . . . . . . . . . . . . . . . . . . 2. Argyreia convolvuloides
1. Bracts very unequal, broadly elliptic, boat-shaped, circular or elliptic, imbricate with calyx, appressed to it;
outer sepals ovate to orbicular or elliptic to ovate, both sides glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Climber; 5 –10 obvious secondary veins; bracts broadly elliptic, boat-shaped, deeply convex; pedicel 1 –5mm
long; outer sepals ovate to orbicular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Argyreia campanuliflora
2. Shrub (rarely twining); indistinct secondary veins; bracts circular or elliptic; pedicel 1 –3 mm long; outer sepals
elliptictoovate...................................................... 3. Argyreia lycioides
Argyreia campanuliflora (Gage)Thoth.(Thothathri
1962: 193).
Lettsomia campanuliflora Gage (Gage 1905: 79). Type:
Burma, Sidoktaya, Minbu distr., Oct. 1902, Shaik
Mokim 418 (lectotype, designated here: CAL!-
image seen [CAL0000018501]; isolectotypes
CAL!-images seen [CAL0000018502,
CAL0000018503, CAL0000018504], K!
[K000197301]).
Blinkworthia convolvuloides sensu Staples & Traiperm
(2017: 471), non Prain.
Climbers, stems 3 –7 m long; branches slender, terete,
base woody, herbaceous above, 8 –10 mm diam.,
brown in vivo.Indumentum simple, non-glandular,
appressed. Leaves with petioles terete, 0.4 –1.8 cm
long, densely hairy; blades oblong-lanceolate to ovate-
lanceolate, 4.2 –14.3 × 1.6 –5.7 cm, chartaceous when
dry, bases usually obtuse or rarely rounded, apices
mucronate-apiculate or acute; hairy on both sides,
adaxially green and darker, sparsely whitish
puberulous, abaxially paler, minutely whitish to yel-
lowish hairy; venation pinnate, secondary veins obvi-
ous, 5 –10 per side, more prominent below, hairs
denser on midvein. Flowers solitary (rarely 1 –3in
cymose inflorescences), axillary, showy, diurnal, pen-
dulous; peduncles 0.6 –2.3 cm long, slender, terete,
densely short-hairy; bracts 4, very unequal, imbricate
with calyx, appressed to it, deeply convex, present at
anthesis, deciduous in fruit, dark green, curved like a
KEW BULLETIN
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boat, adaxially glabrous, margins paler, thinner; outer
2 bracts broadly elliptic, 4 –6×3–5 mm, bases
cuneate, apices obtuse to rounded, abaxially crispate
whitish-hairy; inner 2 bracts boat-shaped (rarely 2-
lobed), 4 –13 × 4 –6 mm, bases attenuate to cuneate,
apexes obtuse, abaxially sparsely whitish-hairy; pedicel
1–5 mm. Sepals unequal, persistent, pale green, both
sides glabrous, margins paler, thinner, adaxially with
several darker veins running straight from base to
apex, outer 2 sepals ovate to orbicular, 11 –22 × 8 –15
mm, margins incurved, inner 3 sepals elliptic-ovate, 10
–16 × 6 –12 mm. Corollas urceolate, 4.7 –4.9 cm long,
waxy-textured in vivo, pale greenish-white outside,
tube inside creamy-white, sparsely magenta-spotted,
limb 5-lobed, recurved, 2.3 –2.7 cm diam. Stamens
included, equal, 15 –20 mm long, white, filaments
dilated basally and thickened at insertion on corolla,
sparsely hairy above insertion, filiform and glabrous
distally; anthers linear-oblong, basifixed, extrorse,
creamy, bases sagittate; pollen globose, spinulose.
Pistils included, usually longer than stamens, 20 –23
mm, white; nectary disc pentagonal, 2 –3 mm long, 5
–6 mm diam., disc surrounding ovary; ovary ovoid,
glabrous c. 5 mm long, c. 3 mm diam., 2-celled, ovules
4, style glabrous, stigmas 2, globose, lobed and
wrinkled. Fruit (immature) loosely enclosed by calyx,
globose, a berry, dark green. Seeds 2–4, c. 12 × 8 mm,
dull black, glabrous. Figs 7;8A, D;9A –B.
DISTRIBUTION. Endemic to the highlands of western
Burma. Map 1.
SPECIMENS EXAMINED.BURMA.Chin State: Mindat, 21
Sept. 1956, F. Kingdon-Ward 22699 (BM!); ibid., alt. 445
–850 m, 4 miles from Kanpetlet and Saw, 29
Aug. 2013, K. Fujikawa et al. 94657 (MBK! image-seen,
QBG!); ibid., 9 Sept. 2002, Ling Shing Maung & Cho
Cho Win 24408 (MBK! image-seen); ibid., alt. 700 –800
m, Natma Taung National Park, 4 Dec. 2002, J. Murata
et al. 24643 (MBK! image-seen); ibid., alt. 900 m, 5
Jan. 2012, S. M. Ling 88122 (MBK! image-seen); ibid., 6
Dec. 2012, K. Fujikawa et al. 90125 (MBK! image-seen);
ibid., alt. 750 m [2400 ft], 10 Dec. 2012, K. Fujikawa
et al. 90187 (MBK! image-seen); ibid., alt. 150 m, 10
Oct. 2012, S. M. Ling 91786 (MBK! image-seen, QBG!);
ibid., alt. 500 m, Kanpetlet Village, 20 Sept. 2016,
P. Rattanakrajang & M. Phyo 113 (BK!, BKF!, QBG!).
Mandalay Region: Myingyan, Wa Laung, 1 Aug. 1909,
J. H. Lace 4882 (E!, K!).
HABITAT. Along roadsides on mountain slopes in
mixed evergreen and deciduous forests; alt. (150 –)
500 –900 m.
CONSERVATION STATUS.Argyreia campanuliflora is en-
demic to western Burma. The extent of occurrence
(EOO) is about 2,184.723 km
2
, below the threshold of
5,000 km
2
for Endangered status. There are two major
populations occuring in at least 12 locations. First, the
population gathered in the urban area, Myingyan
(J. H. Lace 4882 in 1909), has perhaps already been
extirpated. Another population, larger than the
former, was collected in Natma Taung National Park
(IUCN Category II) and nearby areas. However, the
known locations near the protected area have
gradually declined due to agricultural activities and
constructing rural transport routes. Argyreia
campanuliflora is, therefore, assessed here as
Endangered EN B1b(i,ii,iii,iv).
PHENOLOGY. Collected in flower from Aug. to Oct.;
collected in fruit from Oct. to Jan.
ETYMOLOGY. The specific epithet refers to the corolla
shape, which resembles that of Campanula L.
(Campanulaceae).
VERNACULAR NAME. Kyat Hin Cho (Burmese, provided
by villagers at the Kanpetlet village).
USES. The leaves are eaten with chicken soup and
applied as a medicinal shampoo (data from Kanpetlet
villagers at Chin State, Burma).
NOTES.Argyreia campanuliflora can be recognised by
the oblong-lanceolate to ovate-lanceolate leaves, boat-
shaped bracts imbricate with calyx, appressed to it,
and ovate to orbicular outer sepals. A specimen of
A. campanuliflora in Royal Botanic Garden, Edinburgh
(E, J. H. Lace 4882) revealed a 1 –3-flowered cymose
inflorescence (Fig. 8A), which differs from the
othertwospeciesthathavesolitaryorpaired
flowers (Fig. 8B –C).
Argyreia convolvuloides (Prain)Rattanakr. & Traiperm,
comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77304115-1
Blinkworthia convolvuloides Prain, J. Asiat. Soc. Bengal, Pt.
2, Nat. Hist. 63: 91 (1894). Type: Burma, Kendat
[Sagaing], 1890, J. C. Prazer s.n. (lectotype,
selected here: K! [K000830832]; isolectotype
CAL!-image seen [CAL0000018522]).
Blinkworthia discostigma Hand.-Mazz. (Handel-Mazzetti
1931: 6). Type: China, Guangxi, border of Yunnan,
W Poseh, Ba Ko Shan, alt. 370 m [1200 ft], 11
Sept. 1928, R. C. Ching 7353 (lectotype, designated
here: NAS!-image seen [NAS00072335]; isolectotypes
NAS!-images seen [NAS00210035, NAS00210034,
NAS00210033], NY!-image seen [00318857], PE!-
image seen [00029575], W!-image seen).
Climbers (rarely erect shrubs), stems 1 –12 m long,
woody; branches slender, terete, base woody, herba-
ceous above, 5 –8mmdiam.,brownin vivo.
Indumentum simple, non-glandular, appressed. Leaves
with petioles terete, 0.1 –0.6 cm long, densely hairy;
blades elliptic-lanceolate, 1.7 –5.6 × 0.6 –2.3 cm,
leathery, chartaceous when dry, bases usually obtuse,
cuneate or rarely round, apexes usually mucronate
KEW BULLETIN
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apiculate, obtuse, emarginate or rarely acute, adaxially
darker green, glabrous to glabrescent or sparsely
whitish puberulous, abaxially paler, densely whitish to
yellowish hairy; venation pinnate, secondary veins
obvious, 4 –6 per side. Flowers solitary (rarely paired),
axillary, showy, diurnal, pendulous; peduncles 0.2 –
1 cm long, slender, terete, densely short-hairy; bracts 3
or 4, ±equal, spatulate, 3.8 –10 × 3 –5 mm, attached
Fig. 7. Argyreia campanuliflora.Aflowering stem; Bpistil; Cstamens; Dopened corolla with stamens; Eadaxial leaf surface; F
abaxial leaf surface. All from P. Rattanakrajang et al. 113 (BKF). DRAWN BY P.INTHACHUB.
KEW BULLETIN
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below calyx, reflexed from it, margins in-rolled,
present at anthesis, deciduous in fruit, dark green,
both sides short whitish-hairy, base obtuse or cuneate,
margins paler, thinner, apex usually acute or rarely
rounded; pedicels 1 –3 mm. Sepals subequal, persis-
tent, pale green, both sides glabrous, margins paler,
thinner, outer 2 sepals elliptic to ovate or obovate, 6 –
13 × 4 –9 mm; inner 3 sepals elliptic-ovate, 5 –10 × 3 –
8 mm. Corollas tubular-urceolate, 1.7 –2.6 cm long,
waxy-textured in vivo, pale yellowish-white outside,
tube inside creamy-white, magenta-spotted, limb 5-
lobed, recurved, 0.9 –1.9 cm diam. Stamens included,
equal, c. 12 mm long, white, filaments dilated basally
and thickened at insertion on corolla, sparsely hairy
above insertion, filiform and glabrous distally; anthers
linear-oblong, basifixed, extrorse, creamy, bases sagit-
tate; pollen globose, spinulose. Pistils included, usually
shorter than stamens, c. 10 mm long, white; nectary
disc pentagonal, c. 1.5 mm long, c. 3 mm diam., disc
surrounding ovary; ovary ovoid, glabrous c. 3 mm
long, 2 mm diam., 2-celled, ovules 4, style glabrous,
stigmas 2, globose, lobed and wrinkled. Fruit partly
enclosed by reflexed calyx, ovoid, c. 12 mm long, c. 10
diam., brown. Seeds 2–4, c. 10 × 6 mm, dull black,
glabrous. Figs 8B, E;9C –D;10.
DISTRIBUTION. Southern China and northern Burma.
Map 1.
SPECIMENS EXAMINED.CHINA.Guangxi: Tiandong, 3
Oct. 1977, K. D. Liang 3-15033 (GXMI! image-seen);
ibid., alt. 110 m, 27 July 1964, Y. G. Yang 3378 (GXMI!
image-seen); Tianyang, Napo Mountain, 10 Dec. 1964,
K. M. Su 2917 (GXMI! image-seen); Tianlin, Lucheng,
9 June 1964, J. Liu 3348 (GXMI!); Tianlin, alt. 620 m,
21 Nov. 1986, Longtan expedition team 1189 (IBK!
image-seen); Jingxi, 30 Nov. 1967, R. C. Fang & S. H.
Huang s.n. (IBSC! image-seen); Longlin, Nalu, 6
Nov. 1957, Z. D. Nan 4865 (IBSC! image-seen); Baise,
9 Sept. 1953, H. X. Qiu 3744 (IBSC! image-seen, PE!
image-seen); Xilin, 2 July 1936, Z. D. Nan 67904 (IBSC!
image-seen); Longlin, Bada, 29 Oct. 1957, Z. Q. Zhang
10628 (IBSC! image-seen); Baise, Wanglaoshan, 18
Oct. 1989, N. D. Hua 2582 (IBSC! image-seen, PE!
image-seen); ibid., Yangshuo to Menghe Road, 18
July 1958, Z. T. Li 601026 (IBSC! image-seen, KUN!
image-seen); Z. D. Nan 4527 (IBSC! image-seen); ibid.,
Yongle, 21 Sept. 1958, S. Q. Zhong 62361 (KUN!-2
sheets image-seen); ibid., Guangxi expedition team 1277
(KUN! image-seen); ibid., 5 Dec. 1955, S. D. Bai 1127
(PE! image-seen); ibid., Baise to Yongle road, 21
Sept. 1958, S. Q. Zhong A62361 (PE! image-seen).
Yunnan: Yuanjiang, alt. 1000 m, 5 Sept. 2011, L.-Jianwu
1011 (QBG!); Honghe, alt. 450 m, 7 Oct. 1991, G. D.
Tao 47415 (HITBC! image-seen); Yuanyang, Nansha,
18 Oct. 1991, G. D. Tao 47576 (HITBC! image-seen);
ibid., alt. 400 –700 m, 31 May 1974, C. D. Lu 1338
Fig. 8. Comparision of inflorescence and bract characters of three Argyreia species. A, D Argyreia campanuliflora;B, E Argyreia
convolvuloides;C, F Argyreia lycioides.A–Cinflorescence with flowers. A3-flowered cymose inflorescence; B, C paired flowers D
–Fbract shape (from left to right: abaxial, lateral, and adaxial views). Dbroadly elliptic or boat-shaped (upper row: outer bracts,
lower row: inner bracts); Espatulate; Fcircular or elliptic (upper row: outer bracts, lower row: inner bracts). Vouchers:
A. campanuliflora from J. H. Lace 4882 (E); A. convolvuloides from Y. Tang 1040 (KUN) & L. Jianwu 1011 (QBG); A. lycioides from
P. Rattanakrajang & P. Traiperm 2 (BKF). DRAWN BY P.RATTANAKRAJANG.
KEW BULLETIN
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(HITBC! image-seen, KUN! image-seen); alt. 300 m,
13 Nov. 1973, D. D. Tao 1542 (HITBC! image-seen,
IBSC! image-seen, KUN!-2 sheets image-seen); alt. 400
m, 9 Nov. 1973, D. D. Tao 1205 (HITBC! image-seen,
KUN!-3 sheets image-seen); alt. 330 –700 m, 31
May 1974, C. D. Lu 1310 (HITBC! image-seen, KUN!
image-seen); 15 Dec. 1981, X. D. Yu & J. S. Yao s.n.
(IBSC! image-seen); Lushui, alt. 1100 m, 31 July 1960,
Y. Y. Hu 60001644 (KUN! image-seen); Jianshui, alt.
450 m, 27 July 1960, Y. Y. Hu 60001943 (KUN! image-
seen); Yuanjiang, alt. 1025 m, 12 July 2012, Yuanjian
team 5304280909 (IMDY! image-seen); ibid., alt. 900 m,
8 Nov. 1958, S. G. Wu 742 (KUN!-2 sheets image-seen);
ibid., alt. 600 m, 8 Sept. 1959, S. J. Pei 13003 (KUN!
image-seen); ibid., alt. 500 –600 m, 23 Oct. 1965, W. Q.
Yin 1703 (KUN!-2 sheets image-seen); Shiping, alt. 800
Fig. 9. A, B Argyreia campanuliflora.Aplant habit and flowers in side view with boat-like bracts (triangle); Bstem (yellow
triangles). C, D Argyreia convolvuloides.Cflowers in side view with recurved leaf-like bracts (triangle); Dplant habit. E, F Argyreia
lycioides.Eflowers in side view with tiny elliptic-shaped bracts (triangle); Fplant habit. PHOTOS:A–B,E–FP.RATTANAKRAJANG;C–D
K.KOTUN.
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m, 7 July 1988, Y. Tang 1040 (KUN! image-seen);
Hongshiyan, alt. 600 m, 7 May 1984, G. D. Tao 37953
(KUN!-2 sheets image-seen); Ganzhuang, alt. 882 m,
26 Oct. 2013, H. Peng & Z. H. D. Chen 131011 (KUN!-2
sheets image-seen); Yunlong, alt. 2500 m, 1 Feb. 1942,
Q. W. Wang 89756 (KUN! image-seen); Manpan, Red
River, A. Henry 11178 (BM!, E!). BURMA.Kachin State:
Sampanago, bank of Irrawaddy, Ruby mines distr., 5
July 1911, J. H. Lace 5313 (E!, K!). Mandalay Region:
Zibingyi to Thondaung, alt. 600 –900 m [2000 –3000
ft], 30 July 1911, J. H. Lace 5382 (E!, K!); Pyin Ou Lwin
[Maymyo], Thondaung, alt. 730 m, 14 Aug. 1929, Ba
Pe 9606 (RAF!); Thondaung, alt. 790 m, 7 Sept. 1919,
A. Rodger 859 (RAF!); Patheingyi, Ye-Takon Mountain,
alt. 100 m, 23 Sept. 2016, P. Rattanakrajang & M. Phyo
116 (BKF!, QBG!); Pyin Ou Lwin [Maymyo], alt. 850
m, 23 Sept. 2016, P. Rattanakrajang & M. Phyo 117
(BKF!, QBG!). Sagaing Region: Alaungdaw Kathapa
National Park, alt. 300 –360 m, 21 Jan. 2005, J. Murata
et al. 32311 (MBK!); Myingyan, 23 Nov. 1890, J. C.
Prazer s.n. (CAL! image-seen, K!).
HABITAT. Thickets, savanna forest and tropical rain
forest; alt. 100 –1100 (–2500) m.
CONSERVATION STATUS. Near Threatened (NT). In
China, the roots and leaves of Argyreia convolvuloides
are used locally as a medicinal herb, so over-collecting
could reduce the number of mature individuals in
natural habitats. Furthermore, some habitats in
urban locations in Burma (such as Patheingyi and
Pyin Ou Lwin) are at risk for deforestation. The
taxon is very likely declining and fragmented, but it
does not currently evaluate as threatened because
it is widely distributed with an EOO greater than
30,000 km
2
.
PHENOLOGY. Collected in flower from May to Dec.;
collected in fruit from Dec. to Feb.
Map 1. Distribution of Argyreia campanuliflora, circles (○); A. convolvuloides, squares (□); and A. lycioides, triangles (△).
KEW BULLETIN
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2022

ETYMOLOGY. The specific epithet refers to the similar-
ity of the plant habit to the genus Convolvulus L.
VERNACULAR NAMES. Bao ye teng (Chinese, Fang &
Staples 1995), Bindweed-like Blinkworthia (English,
Quattrocchi 2012), Paung khaung long (Burmese, Wai
& Aye 2015), Pan khaung long (Burmese, Maw & Win
2020).
USES. In China the roots and leaves are used for
treating stomachache (Fang & Staples 1995).
NOTES. The name Blinkworthia discostigma Hand.-Mazz.
was widely used on specimens in Chinese herbaria
until it was synonymised under B. convolvuloides by
Fang & Huang (1979). With regard to typification of
B. convolvuloides, there are five sheets filed in two
Fig. 10. Argyreia convolvuloides.Aflowering stem; Bopened corolla with stamens; Cadaxial leaf surface; Dabaxial leaf surface. All
from L. Jianwu 1011 (QBG). DRAWN BY N.CHITCHAK.
KEW BULLETIN
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2022

herbaria, two sheets in Kew (K) and three sheets in
Calcutta Botanic Garden (CAL), all collected by J. C.
Prazer s.n. but coming from two locations in Burma
(Kendat and Myingyan) both localities referred to by
Prain (1894); these five sheets we consider to be syntypes
for this name. The material selected as lectotype best
matches the protologue with complete flowers, the
distinctive leaf-like bracts, hairy on both sides and
attached at the middle between pedicel and peduncle.
Furthermore, the lectotype sheet bears Prain’s handwrit-
ing. The other sheets under J. C. Prazer s.n. from Kendat
are also considered as isolectotypes.
Fig. 11. Argyreia lycioides.Aflowering stem; Bimmature fruit (side view); Cdried fruit (perspective view); Dstamen; Epistil; F
opened corolla with stamens. All from P. Rattanakrajang & P. Traiperm 6 (BKF). DRAWN BY N.CHITCHAK.
KEW BULLETIN
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2022

Argyreia lycioides (Choisy)Traiperm & Rattanakr.,
comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77304116-1
Blinkworthia lycioides Choisy, Mém. Soc. Phys. Genève 6:
430 (1833 publ. 1834). Type: Burma, Kyauk-
Tolong, “Ripa Irawaddi”, 28 Sept. 1826,
N. Wallich 553 [Wall. Cat. 1390] (lectotype,
selected here: G-DC!-image seen [G00135946, 2
sheets]; isolectotypes BM! [BM001011555], G!
[G00227129], K! [K000830830], K-W!
[K001113003]).
Shrubs (rarely twining); stems 0.5 –3 m long, woody;
branches slender, terete, base woody, herbaceous above, 5
–7mmdiam.,brownin vivo.Indumentum simple, non-
glandular, appressed. Leaves with petioles terete, 0.2 –
0.6 cm long, densely hairy; blades elliptic-lanceolate, 0.8
–4.3 × 0.4 –1.5 cm, chartaceous when dry, bases obtuse or
cuneate, apices usually mucronate-apiculate or emargin-
ate, hairy on both sides, adaxially darker green, sparsely
whitish puberulous, abaxially paler, densely whitish to
yellowish hairy; venation pinnate, secondary veins indis-
tinct. Flowers solitary (rarely paired), axillary, showy,
diurnal, pendulous; peduncles 0.3 –1.1 cm long, slender,
terete, densely short-hairy; bracts 4, very unequal, imbri-
cate with calyx, appressed to it, persistent, green, adaxially
glabrous, margins paler, thinner, outer 2 bracts circular or
elliptic, 1 –2×1–2 mm, bases obtuse, minutely whitish-
hairy, apexes rounded; inner 2 bracts elliptic, 3 –5×1–2
mm, bases obtuse, apexes rounded, abaxially glabrous;
pedicels 1 –3mm.Sepals unequal, persistent, pale green,
both sides glabrous, margins paler, thinner; outer 2 sepals
elliptic to ovate, 3 –10 × 3 –7 mm; inner 3 sepals elliptic-
ovate, 2 –8×2–6 mm. Corollas urceolate, 1.1 –2.4 cm
long, waxy-textured in vivo, pale yellowish-white outside,
tube inside creamy-white, magenta-spotted, limb 5-lobed,
recurved, 0.5 –1.9 cm diam. Stamens included, equal, 8 –
10 mm long, white, filaments dilated basally and thick-
ened at insertion on corolla, minutely ciliate above
insertion, filiform and glabrous distally; anthers linear-
oblong, basifixed, extrorse, creamy, bases sagittate; pollen
globose, spinulose. Pistils included, usually longer than
stamens, c. 12 mm long, white; nectary disc pentagonal, c.
2 mm long, c. 4 mm diam., disc surrounding ovary; ovary
ovoid, c. 3 mm long, 2 mm diam., glabrous, 2-celled,
ovules 4, style glabrous, stigmas 2, globose, lobed and
wrinkled. Fruit partly enclosed by reflexed calyx, ovoid, 8 –
11 mm long, 6 –7 mm diam., dull brown. Seeds 2–4, c. 8 ×
5 mm, dull black, glabrous. Figs 8C, F;9E –F;11.
DISTRIBUTION. Burma and northern, eastern to western
Thailand. Map 1.
SPECIMENS EXAMINED.BURMA.Bago Region: Irrawaddy
and Sittang Valley, Feb. 1894, S. Kurz 1085 (K!).
Magway Division: Yenangyaung, Sept. 1934, D. E.
Barnard SSW26 (BM!). Mandalay Division: Meiktila, 1
July 1888, C. B. Collett 867 (K!, L!); Pyinmana, 19
Aug. 1890, Abdul Huk. s.n. (BM!, P!); Popa Village to
Pagan (Janhle Village), 21 Aug. 2000, T. Than Aye &
K. Myo Htwe 20558 (MBK! image-seen); Popa Moun-
tain Park, Kyaukpaduang Township, 9 Oct. 2000,
T. Than Aye & K. Myo Htwe 21175 (MBK! image-seen);
Lein Daw Reserve, Meiktila, 16 Sept. 1969, U. Mg Gale
14635 (RAF!); Yamethin distr., Siulhe Reserve, alt. 200
m [700 ft], 2 Oct. 1909, J. H. Lace 4948 (E!, K!);
Naypyidaw, Yezin, near the Forest Research Institute,
alt. 150 m, 31 May 2016, P. Rattanakrajang et al. 101
(QBG!). Shan State: near the cave at Kinta Dam,
Panlaung and Padalin Cave Wildlife Sanctuary,
Taunggyi, 7 June 2016, Y. Baba et al. 104920 (MBK!
image-seen, QBG!, RAF!). THAILAND.Chiang Mai:
Queen Sirikit Botanic Garden, Mae Rim, alt. 700 m,
27 Aug. 1997, W. Na Nakorn et al. 9875 (BKF!, QBG!);
Doi Inthanon National Park, alt. 450 m, 18 July 1988,
C. Phengklai et al. 6577 (BKF!, K!); ibid., alt. 450 m, 30
July 1988, H. Koyama T-61619 (BKF!, L!); ibid., alt. 450
m, 30 July 1988, N. Fukuoka T-62368 (BKF!, L!); Klang
Waterfall, alt. 400 –450 m, 18 July 1988, N. Fukuoka
T-62061 (BKF!); Hae Doop Reservoir, Doi Tho, alt.
450 m, 23 Oct. 1987, J. F. Maxwell 87-1261 (BKF!, L!);
Doi Sutep-Pui National Park, alt. 1300 m, 23 May 1909,
A. F. G. Kerr 643 (BM!, K!, L!, P!); ibid., 18 Nov. 1910,
A. F. G. Kerr s.n. (BM!); ibid., alt. 650 m, 20 July 1958,
Th. Sørensen et al. 4287 (BKF!, L!); ibid., alt. 450 m, 12
Dec. 1987, J. F. Maxwell 87-1585 (BKF!, L!); ibid., alt.
460 m, 16 Nov. 2014, P. Rattanakrajang & P. Traiperm
14 (BK!, BKF!, QBG!); Mae Rim, alt. 400 m, 7
Sept. 1989, R. Pooma 330 (BKF!); Mae Sa, alt. 800 m,
13 Sept. 1997, T. Santisuk s.n. (BKF!); Mai Klang
Waterfall, 11 Nov. 1965, P. Sangkhachand 37 (BK!);
near Pang Chang, QSBG, Mae Rim, 1 Oct. 1997, BGO
Staff 9690 (QBG!); Wat Phrabat Kuang Pao, Mae
Taeng, 30 Oct. 2010, Staples et al. 1457 (QBG!); Mae
Taeng, alt. 400 m, 6 Oct. 1994, W. Na Nakorn et al. 2349
(QBG!); Mae Thang, alt. 900 m, 21 Aug. 1994, W. Na
Nakorn et al. 1436 (QBG!); Wang Bua Ban, Doi Sutep
National Park, 5 Aug. 1996, W. Na Nakorn et al. 6998
(QBG!); Chiang Dao, alt. 400 m, 1 Dec. 1957, van
Steenis 19631 (L!); Jaum Tong, Mae Soi Valley, alt. 500
m, 8 Sept. 1990, J. F. Maxwell 90-951 (L!); Mae Rim,
Khilek, alt. 400 m, 1 Nov. 2014, P. Rattanakrajang & P.
Traiperm 6 (BK!, BKF!, QBG!); Mae Tang, alt. 350 m, 3
Sept. 1958, Th. Sørensen et al. 4715 (K!). Chiang Rai:
Muang Pajao, 11 July 1931, Put 3965 (BK!, BM!, E!, K!,
L!, P!). Kanchanaburi: Sri Sawat, alt. 150 m, 16 Nov. 1971,
C. F. van Beusekom et al. 3818 (BKF!, K!, L!, P!); ibid., near
Srinakarin Dam, alt. 230 –300 m, 7 Aug. 1982, T. Shimizu
et al. T-28466 (BKF!); ibid., alt. 360 m, 7 Oct. 2014,
P. Rattanakrajang & P. Traiperm 5(BK!,BKF!,QBG!);
Kao Taung, alt. 1100 m, 19 Oct. 1931, A. F. G. Kerr 19764
(BK!, BM!, E!, K!, P!); Saiyok, Namtok (Station), 15
Aug. 1967, Karem 582 (BK!); ibid., alt. 150 m, 3 Sept. 1931,
KEW BULLETIN
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2022

A. F. G. Kerr 19764A (BK!, BM!, K!); Erawan, alt. 100 –
300 m, 2 July 1974, K. Larsen & S. S. Larsen 34012 (K!);
alt. 30 m, Jan. 1918, Winit 534 (K!). Lamphun: Doi Khun
Dahn National Park, 4 Sept. 1967, M. Tagawa et al. 9165
(BKF!); ibid., alt. 825 m, 24 Sept. 1993, J. F. Maxwell 95-
1082 (BKF!, L!); ibid., alt. 850 m, 22 Oct. 1993, J. F.
Maxwell 93-1239 (BKF!, L!); Sriwichai, Doi Mai Takian,
alt. 675 m, 12 Aug. 1996, J. F. Maxwell 96-1077 (BKF!, L!);
See Boh Bahn, 27 Oct. 2010, G. Staples et al. 1448 (BKF!,
QBG!); Mae Tha, Ban Pa Tong, alt. 425 m, 23 Aug. 2010,
M. Norsaengsri 7160 (QBG!); Sraiu, alt. 450 m [1500 ft],
16 July 1915, Winit 404 (E!, K!). Lop Buri: Chai Badan,
BanKlang,20Aug.1975,T. Smitinand 113610 (BKF!,
BM!). Nakhon Sawan: Hua Wai, 28 Aug. 1931, Put 4075
(BK!, BM!, E!, K!, L!, P!). Phetchaburi: Cha-Am, Khao
Sawoei Kapi, 19 Aug. 1995, P. Trisarasri 3(BCU!);ibid.,
Huay Sai, 24 Aug. 1999, C. Niyomdham 5777 (BKF!); ibid.,
22 Sept. 1999, P. Puudjaa & S. Cholkulchana 702 (BKF!);
ibid., near Khao Sawoei Kapi, alt. 40 m, 7 Sept. 2014,
P. Rattanakrajang & P. Traiperm 2(BK!,BKF!,QBG!).
Phrae: Mar Thang Dam, alt. 250 m, 26 June 2012,
M. Norsaengsri & N. Tathana 9591 (BKF!, QBG!).
Ratchaburi: Suan Pueng, alt. 170 m, 1 Oct. 2015,
P. Rattanakrajang & P. Traiperm 49 (BK!, BKF!, QBG!).
Sa Kaeo: Watthana Nakorn, alt. 10 m, 15 Sept. 2018,
P. Traiperm et al. 633 (BKF!). Sukhothai: 25 Aug. 1961,
C. Chermsiriwattana 71 (BK!). Tak: Sam-Ngao, 26
Nov. 1995, R. Pooma 1302 (BKF!); Lan Sang National
Park, alt. 700 m, 17 April 1985, C. Niyomdham 890 (BKF!,
K!, L!, P!);alt. 100 m, 5 Dec. 1957, T. Smitinand 3905 (L!).
Uttaradit: Road to Nam Paad, alt. 500 m, 16 Aug. 2000,
P. Suksathan 2736 (QBG!).
HABITAT. Mixed or lowland dipterocarp or evergreen
forests, usually on limestone; alt. (10 –) 100 –1300 m.
CONSERVATION STATUS. Near Threatened (NT). Argyreia
lycioides is widely distributed throughout Burma and
Thailand. The taxon is likely declining given widespead
habitat disturbance and its collection locations are now
fragmented, in several regions of both countries. Much
of the species’former habitat has been impacted by
forest destruction caused by agricultural and urban
expansion; thus, the number of subpopulations is
decreasing. However, A. lycioides is not currently assessed
as threatened, as it has an EOO greater than 30,000 km
2
.
There are several recent collections in suitable habitats
still remaining in protected areas such as Doi Suthep-Pui
National Park (IUCN Category II) and Panlaung and
Padalin Cave Wildlife Sanctuary (IUCN Category IV).
PHENOLOGY. Collected in flower from April to Nov.;
collected in fruit from April to June and Aug. to Feb.
ETYMOLOGY. The specific epithet refers to the habit of
the plant, which is similar to the genus Lycium L. of the
nightshade family (Solanaceae).
VERNACULAR NAMES. Pan khaung laung, Taungbo-
lelin, Taungbo-lulin (Burmese, Kress et al.2003); Hing
Hai (Thai, Chiang Mai; Smitinand 2014), Mung Kratai
(Thai, Lop Buri & Ratchaburi; Smitinand 2014),
Rakhang Changphueak (Thai, Sukhothai; Smitinand
2014), Rakang Khao (Thai, Lop Buri; Smitinand
2014), Sum Kratai (Thai, Tak; Smitinand 2014), Ya
Chin (Thai, Petchaburi; Smitinand 2014).
NOTES.Argyreia lycioides is unique in Blinkworthia-like
Argyreia species in having tiny flat bracts imbricated with
the calyx at the apex of the pedicel near the receptacle.
Regarding the typification, Choisy clearly had a
specimen with a flower because plate 5 in the protologue
shows the whole flower and dissections with internal floral
details. However, the sheet preserved in the de Candolle
herbarium of the Conservatoire et Jardin Botaniques
Genève (G-DC) bearing Choisy’s handwriting now has no
flowers attached; the second sheet with the same bar code
number [G00135946] has flowers inside the fragment
capsule. We are therefore designating both sheets as the
lectotype in order to encompass complete material.
Acknowledgements
The authors would like to sincerely thank all members of
the Burma Flora Project team —Wattana Tanming,
Yumiko Baba, Jonathan Peeraman, Aye Mya Mon, and
colleagues, of the Kochi Prefectural Makino Botanical
Garden (Japan), the Forest Research Institute (Burma),
and the Queen Sirikit Botanic Garden (Thailand) —for
allowing PR to join their collecting excursion in Burma
in 2016. In addition, Dr Yotsawate Sirichamorn and
Watchara Arthan instructed PR in the molecular phylo-
genetic approaches. We wish to acknowledge the
curators and staff at the herbaria BCU, BK, BKF, BM,
CAL, E, G, GXMI, HITBC, IBK, IBSC, K, KUN, L, NAS,
NY, P, PE, and W for facilitating herbarium specimen
study. PR would also like to thank the Science Achieve-
ment Scholarship of Thailand (SAST) for research
funding and financial support for this study. PT thanks
the Frontier Research Grant, and Mahidol University,
for research funding. Gretchen Wade, Harvard Botany
Libraries, located and obtained critical literature. Heimo
Rainer (W) and Avishek Bhattacharjee (CAL) provided
digital images by permissions of the Directors,
Naturhisorisches Museum Wien of Austria (W) and
Botanical Survey of India (CAL), respectively, for a type
specimen critical to making a lectotype choice. GS would
like to thank Faculty of Graduate Studies, Mahidol
University, for funds that enabled a visit to Mahidol
University to complete this manuscript. We thank the
reviewers for helping improve our manuscript.
Declarations
Conflict of interest. The authors declare that they
have no conflict of interest.
KEW BULLETIN
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2022

Appendix 1. Accession details of samples used
in phylogenetic analyses in this study.
Taxon name, source or geographic region, voucher
with collector number (acronym of herbarium),
GenBank accession number for ITS, trnL-trnF,matK
and rps16 respectively. An asterisk (*) indicates
sequences newly performed in this study; an en-dash
(–) indicates missing data.
IPOMOEEAE:*Argyreia barbigera Choisy:Burma,L. S.
Mang 96014 (BKF), OM699030, OM759910, OM759935,
OM759960. Argyreia campanuliflora (Gage)Thoth.*(1):
Burma, K. Fujikawa et al. 90125 (MBK), OM699023,
OM759903, OM759928, OM759953; *(2): Burma,
P. Rattanakrajang et al. 113 (QBG), OM699024,
OM759904, OM759929, OM759954. Argyreia
capitiformis (Poir.)Ooststr. *(1): Thailand, Mae Hong
Son, P. Rattanakrajang et al. 71 (BKF), OM699033,
OM759913, OM759938, OM759963; *(2): Thailand,
Chiang Mai, P. Rattanakrajang et al. 80 (BKF),
OM699034, OM759914, OM759939, OM759964.
Argyreia caudata Ooststr.: Malaysia, see Simões et al.
(2015), SAN 152877 (SAN), KP261908, KP236602,
KR024892, KR025035. *Argyreia confusa (Prain)
Raizada: Thailand, Staples et al. 1432 (BKF), OM699032,
OM759912, OM759937, OM759962; Argyreia
(Blinkworthia)convolvuloides (Prain)Rattanakrajang &
Traiperm *(1): China, Li Jianwu 1011 (QBG), OM699025,
OM759905, OM759930, OM759955; *(2): Burma, Man-
dalay, P. Rattanakrajang et al. 116 (QBG), OM699027,
OM759907, OM759932, OM759957; *(3): Burma, Pyin
Oo Lwin, P. Rattanakrajang et al. 117 (QBG), OM699026,
OM759906, OM759931, OM759956. Argyreia erinacea
Ooststr.: Malaysia, see Simões et al. (2015), SAN 152883
(SAN), KP261909, KP236603, KR024893, KR025036.
Argyreia henryi Craib: Thailand, see Simões et al.
(2015), Staples 1324 (SING), KP261910, KP236605,
KR024895, KR025038. Argyreia laotica Gagnep.: Thai-
land, see Simões et al. (2015), Staples 1390 (SING),
KP261911, KP236606, KR024896, KR025039. Argyreia
(Blinkworthia)lycioides (Choisy)Traiperm & Rattanakrajang
*(1): Thailand, Kanchanaburi, P. Rattanakrajang & P.
Traiperm 5 (BKF), OM699020, OM759900, OM759925,
OM759950; *(2): Thailand, Chiang Mai, P. Rattanakrajang
&P.Traiperm6 (BKF), OM699017, OM759897,
OM759922, OM759947; *(3): Thailand, Chiang Mai,
P. Rattanakrajang & P. Traiperm 14 (BKF), OM699018,
OM759898, OM759923, OM759948; *(4): Thailand,
Ratchaburi, P. Rattanakrajang & P. Traiperm 49 (BKF),
OM699021, OM759901, OM759926, OM759951; *(5):
Burma, Shan State, Y. Baba et al. 104920 (MBK),
OM699022, OM759902, OM759927, OM759952; *(6):
Burma, Mandalay Region, P. Rattanakrajang & P. Traiperm
101 (QBG), OM699019, OM759899, OM759924,
OM759949. *Argyreia melvillei (S.Moore)Staples:Thailand,
Staples et al. 1419 (BKF), OM699031, OM759911,
OM759936, OM759961. Argyreia mollis (Burm. f.)Bojer:
Thailand, see Simões et al. (2015), Staples 1391 (SING),
KP261912, KP236607, KR024897, KR025040. Argyreia
nervosa (Burm. f.)Bojer (1): Singapore, Singapore Botanic
Garden, Singapore (cultivation), see Simões et al. (2015),
SBG 20031385 (SING), KP261913, KP236608, KR024898,
KR025041; *(2): Thailand, Bangkok, P. Rattanakrajang &
P. Traiperm 141 (BKF), OM699039, OM759919,
OM759944, OM759969. *Argyreia roxburghii (Sweet)
Choisy (1): Thailand, P. Traiperm 482 (BKF), OM699037,
OM759917, OM759942, OM759967; (2): Burma,
K. Fujikawa et al. 95334 (MBK), OM699038, OM759918,
OM759943, OM759968. Argyreia siamensis (Kerr)Staples:
Thailand, see Simões et al. (2015), Staples 1412 (SING),
KP261914, KP236609, KR024899, KR025042. *Argyreia
stenophylla (Kerr)Staples & Traiperm: Thailand,
P. Traiperm 610 (BKF), OM699029, OM759909,
OM759934, OM759959. *Argyreia variabilis Traiperm &
Staples: Thailand, S. Suddee s.n. (BKF), OM699028,
OM759908, OM759933, OM759958. Argyreia zeylanica
(Gaertn.)Voigt *(1): Sri Lanka, P. Traiperm 419 (BKF),
OM699035, OM759915, OM759940, OM759965; *(2): Sri
Lanka, P. Traiperm 420 (BKF), OM699036, OM759916,
OM759941, OM759966. Astripomoea grantii (Rendle)
Verdc.: Tanzania, see Stefanovićet al. (2002), Koyombo
1000 (MO), –, AY101073,–,–.Astripomoea malvacea
(Klotzsch)A.Meeuse: Tanzania, see Stefanović