$¢PhytoKeys

PhytoKeys 259: 229-237 (2025)
DOI: 10.3897/phytokeys.259.157867

Short Communication

Lappula monocarpa, a new synonym of Lappula tenuis

(Boraginaceae)

Dan-Hui Liu!:2°®

1 State Key Laboratory of Ecological Safety and Sustainable Development in Arid Lands, Xinjiang Institute of Ecology and Geography, Chinese Academy of

Sciences, Urumqi, 830011, Xinjiang, China

2 Xinjiang Key Lab of Conservation and Utilization of Plant Gene Resources, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences,

Urumqi 830011, Xinjiang, China

3 Xinjiang Key Laboratory of Biodiversity Conservation and Application in Arid Lands, Xinjiang Institute of Ecology and Geography, Chinese Academy of

Sciences, Urumqi, 830011, Xinjiang, China

Corresponding author: Dan-Hui Liu (liudanhui@ms.xjb.ac.cn)

OPEN Qaceess

Academic editor: Bo Li
Received: 6 May 2025
Accepted: 13 June 2025
Published: 4 July 2025

Citation: Liu D-H (2025) Lappula
monocarpa, a new synonym of
Lappula tenuis (Boraginaceae).
PhytoKeys 259: 229-237. https://doi.
org/10.3897/phytokeys.259.157867

Copyright: © Dan-Hui Liu.
This is an open access article distributed under
terms of the Creative Commons Attribution

License (Attribution 4.0 International - CC BY 4.0).

Abstract

Lappula monocarpa is currently the sole described species within the genus Lappula
characterized by the development of a single nutlet per fruit. However, since its initial
publication, no additional specimens of L. monocarpa have been collected. Critical ex-
amination of type specimens, combined with field surveys and newly collected material,
reveals that the diagnostic trait of L. monocarpa—the presence of a single nutlet per
fruit—is not consistently observed and likely represents an aberrant specimen. More-
over, all other morphological characteristics of L. monocarpa, including indumentum,
corolla, and nutlet features, align fully with those of L. tenuis. Based on this evidence,
L. monocarpa is herein reduced to a synonym of L. tenuis.

Key words: Boraginaceae, Lappula, morphology, new synonym

Introduction

Lappula Moench is one of the largest genera within the tribe Rochelieae of
the family Boraginaceae (Chacon et al. 2016; Vasile et al. 2025). It compris-
es approximately 50 to 80 species distributed across Eurasia, North Africa,
North and South America, and Australia, with a center of diversity located in
Central Asia (Wang 1981; Ovezinnikova 2005, 2021; Weigend et al. 2016).
Members of Lappula are characterized by blue or white corollas with five
throat appendages, a subulate gynobase, and heteromorphic or homomor-
phic nutlets bearing marginal glochids or wings (Popov 1953; Riedl 1967;
Wang 1981; Ovezinnikova 2005). In China, 36 species of Lappula have been
recorded (Zhu et al. 1995), of which at least nine are endemic.

Lappula monocarpa C.J.Wang is a Chinese endemic species, restricted to
Xinjiang Province (Wang 1981). It was originally described based on a spec-
imen collected from Heishantou (Black Mountain), Jimunai County, Xinjiang.
The species is distinguished by the production of a single, horizontally orient-
ed nutlet per fruit, with a disc margin bearing a single row of glochids. In the

229

Dan-Hui Liu: Lappula monocarpa, a new synonym

protologue, Wang (1981) emphasized that these morphological traits are highly
distinctive within Lappula, particularly the presence of a solitary nutlet, which
clearly separates L. monocarpa from all other taxa of Lappula. Concurrent with
the species description, Wang (1981) also established a new monotypic sub-
section, Lappula subsect. Monocarpae C.J.Wang, to accommodate it. However,
since its original description, no additional specimens of L. monocarpa have
been collected beyond the type specimen.

The nutlets of L. monocarpa bear a single row of marginal glochids with
non-connate bases. Based on these traits, Ovczinnikova (2005) assigned
L. monocarpa to ser. Strictae, a classification that contradicts Wang's (1981)
earlier taxonomic treatment. Subsequently, Ovezinnikova (2009) proposed that
L. monocarpa may represent only a variant of L. stricta, distinguished solely by its
production of a single nutlet per fruit. Nutlet characteristics are extensively uti-
lized for infrageneric classification and species delineation within Lappula. These
characteristics include the presence of glochids or wings, the number of glochid
rows, the length of the glochids, the width of the wings, and the shape of the dor-
sal disk (Giirke 1894; Popov 1953; Riedl 1967; Wang 1981; Ovezinnikova 2005,
2009, 2021). However, the number of nutlets per fruit has not been employed as
a diagnostic character for species delineation, as all currently described species
of Lappula typically develop four nutlets per fruit, with the exception of L. mono-
carpa (Popov 1953; Wang 1981; Ovezinnikova 2005, 2021, 2023; Liu et al. 2024).

Further examination of the type specimen of L. monocarpa revealed that it bears
only a few developing fruits, each containing a single nutlet. This nutlet is ovoid
and features a single row of glochids (Fig. 1). Apart from the difference in nutlet
number, all other nutlet characteristics align fully with those of L. tenuis (Ledeb.)
Gurke. These observations suggest that L. monocarpa and L. tenuis represent the
same taxonomic entity. To test this hypothesis, field surveys were conducted at
the type locality of L. monocarpa, and additional specimens were collected. Mor-
phological comparisons among L. monocarpa, L. tenuis, and L. stricta (Ledeb.)
Gurke were then carried out to clarify the taxonomic status of L. monocarpa.

Material and methods

Herbarium specimens of Lappula from ALTB, BNU, HNWP, KUZ, LE, and NSK
were thoroughly examined, including the type specimens of L. monocarpa and
L. tenuis. A field survey was conducted at the type locality of L. monocarpa in
Heishantou (Black Mountain), Jimunai County, Xinjiang Province, China, where
additional specimens were collected and deposited at BNU. Detailed morpho-
logical features, including stem indumentum, calyx, corolla, nutlets, and gyno-
base, were documented in situ using a Sony Alpha 7 camera. Comparative mor-
phological analyses of L. monocarpa, L. stricta, and L. tenuis were performed to
assess their taxonomic boundaries.

Results

During field investigations, no Lappula specimens matching the original descrip-
tion of producing a single nutlet per fruit were encountered. All collected individu-
als consistently developed four nutlets per fruit. Aside from this numerical discrep-
ancy, their morphological features were indistinguishable from those described for

PhytoKeys 259: 229-237 (2025), DOI: 10.3897/phytokeys.259.157867 930

Dan-Hui Liu: Lappula monocarpa, a new synonym

Figure 1. Type specimen of Lappula monocarpa C.J.Wang. A. Holotype in the herbarium of the Northwest Institute of
Plateau Biology, Chinese Academy of Sciences (HNWP); B, C. Details of the holotype; B. Development of a single nutlet
per fruit, with the nutlet horizontally oriented; C. Nutlet morphology.

PhytoKeys 259: 229-237 (2025), DOI: 10.3897/phytokeys.259.157867

Dan-Hui Liu: Lappula monocarpa, a new synonym

L. monocarpa. Detailed morphological comparisons among L. monocarpa, L. stric-
ta, and L. tenuis revealed congruent characteristics in stem indumentum, bract
morphology, calyx, and corolla (Fig. 2, Table 1). The primary diagnostic differences
were confined to nutlet morphology: both L. monocarpa and L. tenuis exhibit ovate
dorsal disks (length-to-width ratios of 2.5—3), with the greatest width occurring be-
low the midpoint. These nutlets lack a disk keel and possess non-thickened mar-
gins. In contrast, nutlets of L. stricta are lanceolate (length-to-width ratio ~4), may
or may not display a disk keel, and typically have thickened margins.

Discussion

Ovezinnikova (2005) circumscribed Lappula sect. Lappula to include five series:
ser. Lappula, ser. Redowskianae Ovczinnikova, ser. Anisacanthae Ovczinnikova,
ser. Strictae M.Pop. ex Ovezinnikova, and ser. Patulae Ovezinnikova (Table 2).
Recently, Liu et al. (2025) reconstructed the phylogenetic relationships of Asian

Figure 2. Morphological features of Lappula monocarpa C.J.Wang (A-J) and nutlet morphology of L. stricta (Ledeb.)
Gurke (K) and L. tenuis (Ledeb.) Gurke (L). A. Characteristics of stem trichomes, showing appressed and spreading
hairs; B. Calyx; C, D, E, F. Flower morphology; C, D. Lateral view of the flower; E. Top view of the flower; F. Expanded flower
morphology, showing the corolla throat appendages and stamens; G. Gynobase morphology; H, I, J. Nutlet morphology;
H. Abaxial view of the nutlet; I. Lateral view of the nutlet; J. Adaxial view of the nutlet; K. Nutlet morphology of L. stricta;
L. Nutlet morphology of L. tenuis. Scale bars: 1 mm.

PhytoKeys 259: 229-237 (2025), DOI: 10.3897/phytokeys.259.157867 932

Dan-Hui Liu: Lappula monocarpa, a new synonym

Table 1. Morphological comparisons of Lappula monocarpa, L. stricta, and L. tenuis.

Characters L. monocarpa L. stricta L. tenuis
Life form annual annual annual
Indumentum spreading and appressed hair spreading and appressed hair spreading and appressed hair
Bracts leaf-like at the base of the leaf-like at the base of the leaf-like at the base of the
inflorescence, gradually reduced inflorescence, gradually reduced inflorescence, gradually reduced
toward the apex toward the apex toward the apex
Calyx lobes linear linear linear
length exceeding the fruit exceeding the fruit exceeding the fruit
Corolla length ca.3 mm ca.3 mm ca.3 mm
Nutlet | glochids single row single row single row
disk shape ovate lanceolate ovate
disk keel without with or without without
margin not thickened thickened not thickened
Style slightly surpassing nutlet slightly surpassing nutlet slightly surpassing nutlet

Table 2. The subdivision of Lappula sect. Lappula by Ovczinnikova (2005), including the species composition of each series.

Taxa

Ser. Lappula
L. brachycentroides
L. consanguinea
L. heteracantha
L. intermedia
L. Squarrosa
L. tenuis
L. tuvinica

Sect. Lappula

Ser. Redowskianae Ser. Anisacanthae Ser. Strictae Ser. Patulae
L. occidentalis L. anisacantha L. caespitosa L. patula
L. redowskii L. lenensis L. coronifera
L. shanhsiensis L. cristata

L. fruticulosa
L. karelinii
L. marginata
L. monocarpa
L. physacantha
L. stricta
L. zaissanica

Lappula using hundreds of single-copy nuclear genes and complete chloroplast
genomes. Their results support the monophyly of sect. Lappula as defined by
Ovezinnikova (2005). However, the placement of certain taxa conflicts with this
classification. Notably, L. monocarpa clustered with L. consanguinea and L. ano-
carpa, indicating a closer affinity with ser. Lappula. Furthermore, L. monocarpa
and L. stricta occupy distinct, strongly supported subclades within sect. Lappu-
la (Clade IV; Liu et al. 2025). This phylogenetic topology supports neither the in-
clusion of L. monocarpa in ser. Strictae (Ovezinnikova 2005), nor the hypothesis
that L. monocarpa represents a variant of L. stricta (Ovezinnikova 2009).
Ovezinnikova (2005) defined ser. Strictae as comprising taxa with nutlets
bearing a single row of glochids, typically with broadened bases or, occasionally,
forming marginal wings. In contrast, ser. Lappula includes nutlets with one to
several rows of glochids, the bases of which are neither broadened nor winged.
Ovezinnikova assigned L. monocarpa to ser. Strictae and L. tenuis to ser. Lappula.
However, both the type specimen of L. monocarpa and newly collected material
from its type locality reveal that the glochids have neither broadened nor winged
bases, and no winged nutlets were observed in the population. These morpho-
logical traits are inconsistent with the defining characteristics of ser. Strictae.
Detailed morphological comparisons between the Asian Lappula taxa L. stric-
ta (ser. Strictae) and L. tenuis (ser. Lappula) reveal that the primary distinguish-

PhytoKeys 259: 229-237 (2025), DOI: 10.3897/phytokeys.259.157867 933

Dan-Hui Liu: Lappula monocarpa, a new synonym

ing feature between these two series lies in the shape of the nutlet disk, rather
than in the number of glochid rows. The “disk” refers to the flattened area on the
dorsal side of the nutlet, the margin of which typically bears glochids or wings
(Hilger 2014). Nutlets of ser. Strictae are lanceolate and may or may not pos-
sess a keel, whereas those of ser. Lappula are ovate and consistently keel-less.
The nutlet disk of L. monocarpa is ovate and lacks a keel, supporting its place-
ment within ser. Lappula on morphological grounds, in agreement with the phy-
logenetic results of Liu et al. (2025). Additionally, some North American Lappula
taxa, such as L. desertorum Greene and L. heterosperma Greene, exhibit a ridge
on the dorsal face of the nutlet or lines of tubercles along the midline of the
disk (Rolfsmeier 2013). Liu et al. (2025) suggested that these North American
taxa may be closely related to Clade IV on the Eurasian continent. The species
examined in the present study, L. monocarpa and L. stricta, also belong to Clade
IV. While sect. Lappula, as defined by Ovezinnikova (2005), includes only one
native North American species, L. occidentalis (S. Watson) Greene, the phyloge-
netic placement of other North American Lappula taxa remains uncertain and
requires further investigation using both molecular and morphological data.
As defined by Ovezinnikova (2005), ser. Lappula comprises seven species,
among which only L. tenuis and L. brachycentroides Popov produce nutlets with
a single row of spines. Popov (1951) noted that L. brachycentroides is closely
related to Echinospermum tenue Ledeb. (now accepted as L. tenuis), differing
primarily by the presence of short or absent glochids on the nutlets. The nutlets
of L. monocarpa bear a single row of glochids with non-broadened bases and ex-
hibit an ovate, non-lanceolate dorsal disk. Taken together, the molecular phyloge-
netic findings of Liu et al. (2025) and the morphological evidence presented here
support the placement of L. monocarpa within ser. Lappula. This classification
supports the reduction of L. monocarpa to a taxonomic synonym of L. tenuis.

Taxonomic treatment

Lappula tenuis (Ledeb.) Giirke, Nat. Pflanzenfam. 4(3a): 107, 1894

= Echinospermum tenue Ledeb, FI. Altaic. 1: 201, 1829.

= Hackelia tenuis (Ledeb.) Opiz, Oekon.-Techn. Fl. Bohm. 2(2): 147, 1839.

= Myosotis tenuis (Ledeb.) Morch, Cat. Hort. Hafn.: 64, 1839.

= Echinospermum redowskii var. tenue (Ledeb.) Regel, Bull. Soc. Imp. Natural-
istes Moscou 41(I): 84, 1868.

= Cynoglossospermum tenue (Ledeb.) Kuntze, Revis. Gen. PI. 2: 437, 1891.

= Lappula monocarpa C.J.Wang, Bull. Bot. Res., Harbin 1(4): 98, 1981. syn. nov.
Type. China, Xinjiang province, Jimunai County, 1130 m a.s.l., 22 August
1974, Xinjiang Exped. 1750 (Holotype: HNWP)).

Type. RussiA « Altai, Ad fluv. Tscharysch, 1826, Ledebour, Smejow and Politow
s. n. (Lecotype by Ovezinnikova (2018), Lectotype: LE01043915!).

Distribution and habitat. Lappula tenuis is distributed in China, Kazakhstan,
and Russia (Wang 1981, Ovezinnikova 2009, Chepinoga et al. 2024). This spe-
cies inhabits grasslands and steppes at altitudes of approximately 1,000 me-
ters above sea level.

Phenology. Flowering and fruiting from June to August.

PhytoKeys 259: 229-237 (2025), DOI: 10.3897/phytokeys.259.157867 934

Dan-Hui Liu: Lappula monocarpa, a new synonym

Taxonomic notes. Fruits of Lappula typically develop four nutlets; however, an
extensive review of herbarium specimens revealed that some individuals, for ex-
ample, L. caespitosa C.J.Wang (BNU0033484) and L. microcarpa (Ledeb.) Giirke
(BNU0030476, BNU0030622, E00843758), sometimes produce only a single
developed nutlet. Due to the infrequent and irregular occurrence of such aber-
rations, nutlet number is considered an unreliable diagnostic character for spe-
cies delimitation within the genus. During field surveys, L. tenuis was occasion-
ally found growing sympatrically with L. consanguinea (Fisch. & C.A.Mey.) Giirke
and L. anocarpa C.J.Wang. Although these three species share similar vegeta-
tive morphology and ovoid nutlet shape, they can be reliably distinguished by
the arrangement of glochids: L. tenuis has nutlets with a single row of marginal
glochids, whereas L. consanguinea and L. anocarpa bear two or three rows.

Ovezinnikova (2005) circumscribed ser. Lappula to include only two species
with single-rowed marginal glochids: L. tenuis and L. brachycentroides. Popov
(1951) recognized the morphological similarity between these taxa but distin-
guished L. brachycentroides by the presence of short or absent glochids along
the nutlet margins. However, my examination of L. brachycentroides specimens
revealed that nutlets on the same individual can exhibit either smooth margins
or distinct glochids, with the latter morphology being consistent with that of
L. tenuis. Fruit heteromorphism is well documented in the genus Lappula, in-
cluding variation in glochid presence (glochids or wings, long-glochids, short-
glochids, or glochidless) and wing morphology (broad- or narrow-winged).
These observations suggest that L. brachycentroides may simply represent a
fruit-dimorphic form of L. tenuis. However, as no newly collected material of
L. brachycentroides was obtained in this study, the taxonomic status of this
species requires further investigation.

Additional specimens examined. CHINA. Xinjiang: +» Altai, Xinjiang Exped.
10669 (PE01361016); * Jimunai County, D.H. Liu BNU2019XJ214 (BNU); + Qin-
ghe County, D.H. Liu BNU2019XJ182 (BNU). KAZAKHSTAN. * Akmola region,
Lashchinsky N.N. s.n. (NSKO0009103); » East Kazakhstan, Kotukhov A.Yu. s.n.
(NSK0008186); » Zharminsky district, Korolyuk A. Yu. 132 (NSK0005792); * Kok-
pekti district, Popov N.V. 7 (NSKO005797); RussIA. « Altai region, Usyk N.A. s.n.
(ALTB1100059605, ALTB1100060899); *» Kemerovo region, Stre/nikova T.O. and
Manakov Y.A. KEM18465 (KUZ026052); * Kosh-Agachsky district, Korolyuk A. Yu.
121 (NSK0005798); * Ongudaysky district, Maslova O.M. and Khrustaleva |.A. s.n.
(NSK0005819); * Orenburg region, Lashchinsky N.N. L15-174 (NSK0009078).

Acknowledgements

The author expresses sincere gratitude to the curators of the herbaria ALTB,
BNU, HNWP, KUZ, LE, and NSK for providing access to specimens and digital
images for examination. Appreciation is also extended to the anonymous re-
viewer for their valuable and constructive comments, which substantially im-
proved the quality of the manuscript.

Additional information
Conflict of interest

The author has declared that no competing interests exist.

PhytoKeys 259: 229-237 (2025), DOI: 10.3897/phytokeys.259.157867 995

Dan-Hui Liu: Lappula monocarpa, a new synonym

Ethical statement

No ethical statement was reported.

Use of Al

No use of Al was reported.

Funding

This work was supported by the Tianchi Talent Introduction Program of the Xinjiang
Uygur Autonomous Region; the Natural Science Foundation of the Xinjiang Uygur Auton-
omous Region (grant no. 2023D01B02); and the Biological Resources Programme of the
Chinese Academy of Sciences (grant no. CAS-TAX-24-070).

Author contributions

Conceptualization: DHL. Data curation: DHL. Formal analysis: DHL. Funding acquisition:
DHL. Writing - original draft: DHL. Writing - review and editing: DHL.

Author ORCIDs
Dan-Hui Liu © https://orcid.org/0000-0002-0195-1436

Data availability

All of the data that support the findings of this study are available in the main text.

References

Chacon J, Luebert F, Hilger HH, Ovchinnikova S, Selvi F, Cecchi L, Guilliams CM, Hasen-
stab-Lehman K, Sutory K, Simpson MG, Weigend M (2016) The borage family (Bor-
aginaceae s.str.): A revised infrafamilial classification based on new phylogenetic
evidence, with emphasis on the placement of some enigmatic genera. Taxon 65:
523-546. https://doi.org/10.12705/653.6

Chepinoga VV, Barkalov VY, Ebel AL, Knyazev MS, Baikov KS, Bobrov AA, Chkalov AV,
Doronkin VM, Efimov PG, Friesen NV, German DA, Gontcharov AA, Grabovskaya-Boro-
dina AE, Gureyeva II, lvanenko YA, Kechaykin AA, Korobkov AA, Korolyuk EA, Kosachev
PA, Kupriyanov AN, Luferov AN, Melnikov DG, Mikhailova MA, Nikiforova OD, Orlova
LV, Ovchinnikova SV, Pinzhenina EA, Poliakova TA, Shekhovstsova IN, Shipunov AB,
Shmakov Al, Smirnov SV, Tkach N, Troshkina VI, Tupitsyna NN, Vasjukov VM, Vlasova
NV, Verkhozina AV, Anenkhonov OA, Efremov AN, Glazunov VA, Khoreva MG, Kiseleva
TI, Krestov PV, Kryukova MV, Kuzmin IV, Lashchinskiy NN, Pospelov IN, Pospelova
EB, Zolotareva NV, Sennikov AN (2024) Checklist of vascular plants of Asian Russia.
Botanica Pacifica 13: 3-310. https://doi.org/10.17581/bp.2024.13s01

Gurke M (1894) Lappula Moench. In: Prantl K, Engler A (Eds), Die natiirlichen Pflanzen-
familien Engelmann, W Engelmann, Leipzig, Vol. 4, 106-107.

Hilger HH (2014) Ontogeny, morphology, and systematic significance of glochidiate and
winged fruits of Cynoglosseae and Eritrichieae (Boraginaceae). Plant Diversity and
Evolution 131(3): 167-214. https://doi.org/10.1127/1869-6155/2014/0131-0080

Liu DH, Zhou YX, Shang SJ, Wu JJ, Li WJ (2024) Lappula effusa (Boraginaceae), a new
species from Xinjiang, China. PhytoKeys 243: 105-112. https://doi.org/10.3897/phy-
tokeys.243.123468

Liu DH, Liu QR, Tojibaev KS, Sukhorukov AP, Wariss HM, Zhao Y, Yang L, Li WJ (2025)
Phylogenomics provides new insight into the phylogeny and diversification of Asian

PhytoKeys 259: 229-237 (2025), DOI: 10.3897/phytokeys.259.157867 036

Dan-Hui Liu: Lappula monocarpa, a new synonym

Lappula (Boraginaceae). Molecular Phylogenetics and Evolution 208: 108361. https://
doi.org/10.1016/j.ympev.2025.108361

Ovezinnikova SV (2005) The system of the subtribe Echinosperminae (tribe Eritrichieae,
Boraginaceae). Botanicheskii Zhurnal 90(8): 1153-1172. [in Russian]

Ovezinnikova SV (2009) The synopsis of the subtribe Echinosperminae Ovczinnikova
(Boraginaceae) in the flora of Eurasia. Novosti sistematiki vysshikh rasteniy 41: 209-
272. [in Russian] https://doi.org/10.31111/novitates/2010.41.209

Ovezinnikova SV (2018) Type specimens of names of Altai taxa of Boraginaceae de-
scribed by C.F. Ledebour, A.A. Bunge, and C.A. Meyer, kept in the Herbarium of Ko-
marov Botanical Institute (LE). Novitates Systematicae Plantarum Vascularium 49:
112-124. [in Russian] https://doi.org/10.31111/novitates/2018.49.112

Ovezinnikova SV (2021) A new species Lappula botschantzevii (Boraginaceae) from
the Northern Africa. Phytotaxa 522(1): 47-55. https://doi.org/10.11646/phyto-
taxa.522.1.5

Ovezinnikova SV (2023) New taxa of the genus Lappula (Boraginaceae) from Eastern
Siberia. Flora and Vegetation of Asian Russia 2: 130-142.

Popov MG (1951) Boraginaceae Novae. In: Shishkin BK (Ed.) Botanicheskie Materialy
Gerbariya Botanicheskogo Instituta Imeni V. L. Komarova Akademii Nauk SSSR, Vol.
14. Moscow, 314-317. [in Russian]

Popov MG (1953) Lappula Moench. In: Komarov VL (Ed.) Flora USSR, Vol. 19. Akademii
Nauk SSSR, Leningrad, 404-479.

Riedl H (1967) Lappula Moench. In: Rechinger KH (Ed.) Flora Iranica, Vol. 48. Akade-
mische Druck- und Verlagsanstalt, Graz, Austria, 68-71.

Rolfsmeier SJ (2013) Taxonomy and phylogeny of the genus Lappula Moench (Borag-
inaceae) in North America. PhD Thesis, Kansas State University, United States.

Vasile MA, Bohnert T, Jeiter J, Cardoso D, Moonlight PW, Weigend M (2025) An updated
phylogeny of Boraginales based on the Angiosperms353 probe set: A roadmap for
understanding morphological evolution. Annals of Botany XX: mcaf061. https://doi.
org/10.1093/aob/mcaf061

Wang CJ (1981) A study on the genus Lappula of China. Bulletin of Botanical Research
1(4): 77-100.

Weigend M, Selvi F, Thomas DC, Hilger HH (2016) Boraginaceae. In: Kadereit JW, Bittrich
V (Eds) The families and genera of vascular plants. Vol. 14. Flowering plants: Eudicots;
Aquifoliales, Boraginales, Bruniales, Dipsacales, Escalloniales, Garryales, Paracryph-
iales, Solanales (except Convolvulaceae), Icacinaceae, Metteniusaceae, Vahliaceae.
Springer International Publishing, Cham, 41-102. https://doi.org/10.1007/978-3-
319-28534-4_5

Zhu GL, Riedl H, Kamelin RV (1995) Lappula Moench. In: Wu ZY, Raven PH, Hong DY
(Eds) Flora of China, Vol. 16. Science Press, Beijing & Missouri Botanical Garden
Press, St. Louis, 402-414.

PhytoKeys 259: 229-237 (2025), DOI: 10.3897/phytokeys.259.157867 997