Lappula effusa (Boraginaceae), a new species from Xinjiang, China

Dan-Hui Liu; Yi-Xin Zhou; Shu-Jing Shang; Jia-Ju Wu; Wen-Jun Li

doi:10.3897/phytokeys.243.123468

Abstract

Lappula effusa D.H.Liu & W.J.Li, a new species of Boraginaceae from Xinjiang, China, is described and illustrated in this study. The new species is morphologically similar to Lappula himalayensis and L. tadshikorum. However, it can be distinguished from the compared species by several characteristics, such as: stem single, erect, frequently branched at middle and above, densely spreading hispid, hairs discoid at base; corolla white or blue; fruit compressed, heteromorphic nutlets with two rows of marginal glochids, nutlets acute ovoid, disc narrowly ovate-triangular. The diagnosis of the new species is supported with comprehensive investigation including photographs, detailed description, notes on etymology, distribution and habitat, conservation status, as well as comparisons with morphologically similar species.

Key words

Boraginaceae, China, Lappula, new taxon, taxonomy

Introduction

The genus Lappula Moench, belonging to the Boraginaceae family within the Rochelieae tribe, encompasses approximately with 50–70 species (Ovczinnikova 2005; Weigend et al. 2016). These species are predominantly distributed in Eurasia, North Africa, North and South America and Australia, with the centre of species diversity lying in Central Asia (Wang 1981; Ovczinnikova 2005, 2021; Huang et al. 2013). Lappula is characterised by prickly cauline leaves, blue/white corollas that each bear five throat appendages, a subulate gynobase, nutlets four, homomorphic or heteromorphic and nutlets with either one/more rows of marginal glochids or marginal wings tipped with anchor-like spines (Popov 1953; Riedl 1967; Zhu et al. 1995; Huang et al. 2013; Weigend et al. 2016; Khoshsokhan-Mozaffar et al. 2018).

Initially, Lappula had been treated as a member of Myosotis L. (Linnaeus 1753), with Moench (1794) later distinguishing and circumscribing Lappula as a separate genus. In the taxonomy of Lappula originating from Candolle (1846), there were 38 species in the Prodromus and these species were classified into three sections, based on the morphology of nutlets. In the Flora USSR, Popov (1953) identified 39 species and improved the infrageneric classification of Lappula by introducing two sections and 14 series. Ovczinnikova (2005) recognised 70 species and proposed an updated infrageneric classification of Lappula, based on corolla, nutlets and gynobase morphology. She classified the 70 species into eight sections and 14 series. Recent molecular phylogenetic studies showed that Lappula was polyphyletic and some species were transferred to the Rochelia and Pseudolappula (Huang et al. 2013; Khoshsokhan-Mozaffar et al. 2018; Liu et al. 2021). In China, the genus Lappula had 31–36 species and was highly diversified in north-western China, especially in Xinjiang Province (Wang 1981; Zhu et al. 1995).

During field investigations in Xinjiang Province, China, an unknown population of Lappula was discovered in Balikun County. It appeared to be similar to L. himalayensis Ching J.Wang and L. tadshikorum Popov in general habit and fruit morphology. However, the unknown population showed great differences in an array of characters: stem single, frequently branched at middle and above, spreading; style surpassing the fruit by ca. 0.5 mm, fruit compressed, nutlets acute ovoid and disc narrowly ovate-triangular (Figs 1, 3). After conducting a comprehensive review of relevant literature (Gürke 1894; Brand 1931; Popov 1953; Vvedensky 1961; Sharashova 1962; Goloskokov 1964; Riedl 1967; Chukavina 1984; Nasir 1989; Zhu et al. 1995; Ovczinnikova 2009) and examining specimens of Lappula from the Herbaria of Royal Botanic Garden Edinburgh (E), Muséum National d’Histoire Naturelle (P), Komarov Botanical Institute of RAS (LE), Moscow University (MW), Central Siberian Botanical Garden SB RAS (NS), Institute of Botany, Chinese Academy of Sciences (PE), North-western Institute of Botany (WUK), Northwest Institute of Plateau Biology, Chinese Academy of Sciences (HNWP), National Herbarium of Uzbekistan (TASH) and Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences (XJBI), we concluded that it did not match morphologically with any known species of Lappula. Based on these distinctive morphological features, we confirmed that it was a new species, which we describe and illustrate here as Lappula effusa D.H.Liu & W.J.Li.

Figure 1.

Lappula effusa D.H.Liu & W.J.Li, sp. nov. A habitat B habit C, D flower morphology E inflorescences F fruit G spreading calyx in fruit.

Materials and methods

The voucher specimens of the new species in this study were collected during our field expedition to Xinjiang Province in 2023. Photographs were captured using a Nikon Z7 II digital camera (Tokyo, Japan), while morphological observations and measurements were conducted on living plants in the field and herbarium specimens deposited at XJBI. For morphological comparison, we critically examined available digitised specimens of Lappula stored in the E (https://data.rbge.org.uk/search/ herbarium/), LE (https://plant.depo.msu.ru/), MW (https://en.herbariumle.ru/), NS (http://herb.csbg.nsc.ru:8081/) and P (https://science.mnhn.fr/institution/mnhn/collection/p/item/search), as well as physical herbarium specimens deposited at PE, HNWP, TASH, WUK and XJBI. Additionally, we compared the morphological characteristics of the new species with those of similar species, relying on online or physical specimens. The conservation status was assessed following the IUCN guidelines (IUCN 2022). In this study, we employed the morphological species concept (Davis and Heywood 1963; Mallet 1995), which defines species solely by their morphological differences.

Results

Taxonomy

Lappula effusa D.H.Liu & W.J.Li, sp. nov.

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Figs 1, 2, 3

Diagnosis

The new species is morphologically similar to Lappula himalayensis and L. tadshikorum, but differs from the L. himalayensis primarily in the following characteristics: stem single (vs. stems 4–6, cespitose), erect (vs. ascending or erect), frequently branched at middle and above (vs. branched above), densely spreading hispid, hairs discoid at base (vs. densely appressed pubescent); corolla white or blue (vs. blue); fruit compressed (vs. fruit globose), nutlets acute ovoid, ca. 2.5 mm long, 1 mm wide, 0.5 mm thickness (vs. ovoid, ca. 2.5 mm long, 1.5 mm wide, 1 mm thickness), the inner glochids erect (vs. often curved), disc narrowly ovate-triangular (vs. ovoid) (Figs 2, 3). Furthermore, compared to the L. tadshikorum, the new species is 12–28 cm tall (vs. 30–50 cm tall), stem single (vs. stems 2–3), densely spreading hair (vs. appressed or semi-appressed hair), fruit compressed (vs. fruit globose-ovoid), nutlets acute ovoid, 0.5 mm thickness (vs. ovoid, 1 mm thickness), disc narrowly ovate-triangular (vs. oblong or ovate), inner glochids erect, ca. 0.5 mm long (vs. curved, 1–1.2 mm long), style surpassing nutlets and glochids (vs. style slightly surpassing nutlets, but not surpassing glochids) (Figs 2, 3).

Figure 2.

Type specimens of L. himalayensis, L. tadshikorum and L. effusa A holotype of L. himalayensis (PE00029615!) B lectotype of L. tadshikorum (LE 140!) C holotype of L. effusa (XJBI00135936!).

Figure 3.

Morphological comparisons of L. himalayensis, L. effusa and L. tadshikorum. L. himalayensis A stem indumentum B gynobase C fruit lateral view D fruit polar view E nutlet abaxial view (with short glochids) F nutlet lateral view (with long glochids) G nutlet lateral view (with short glochids) H nutlet adaxial view. L. effusa I stem indumentum J gynobase K fruit lateral view L fruit polar view M nutlet abaxial view (with short glochids) N nutlet abaxial view (with long glochids) O nutlet lateral view P nutlet adaxial view. L. tadshikorum Q stem indumentum R gynobase S fruit lateral view T fruit polar view U nutlet abaxial view (with short glochids) V nutlet abaxial view (with long glochids) W nutlet lateral view X nutlet adaxial view. Scale bar represents 0.5 mm.

Type

China. Xinjiang: Balikun County, Dahongliuxia Village, growing on the gravel desert, 44°47'26.17N, 91°30'9.55E, alt. 842 m, 18 June 2023, D.H.Liu, Y.X.Zhou, S.J.Shang et al. 2023EH908 (holotype: XJBI00135936!).

Description

Annual herbs. Stems erect, single, frequently branched at middle and above, 12–28 cm tall, with spreading white hispid, hairs discoid at base (Fig. 3I). Basal leaves forming a rosette; leaf blade spatulate, 1.5–2.5 cm long, 2–4 mm wide, densely spreading white hirsute, hairs discoid at base; withered in fruit; Stem leaves linear-lanceolate, 1–2 cm long, 2–4 mm wide, abaxially densely spreading white hispid, adaxially sparsely hispid or glabrous, hairs discoid at base. Inflorescences elongated to 5–10 cm long in fruit, with oval bracts near 3–5 mm long, 1.5–2.5 mm wide. Pedicels short, ca. 1 mm long in flowering and elongated from 2.5–3 mm long in fruit. Calyx lobes oblong, ca. 1.5 mm long, 1 mm wide, slightly elongated ca. 2 mm long in fruit, spread, abaxially densely spreading hispid, adaxially sparsely hispid or glabrous. Corolla blue and white (plants with either all blue or all white corollas), campanulate, corolla tube ca. 1.5 mm long; limb as long as tube, ca. 1.5 mm wide, lobes obtuse; throat appendages white or light yellow, trapeziform, ca. 0.3 mm high; stamens five, included in the corolla tube, filament short, inserted at the middle of tube, anthers brown. Gynobase narrowly subulate (Fig. 3J), with a style surpassing the nutlets by ca. 0.5 mm. Coenobium laterally compressed ovoid (Fig. 3L), with glochids 2.5–3 mm in diameter. Nutlets four, heteromorphic, easily separated from gynobase, acute ovoid; 2.5–3 mm long, ca. 1.2 mm wide, disc narrowly ovate, adaxially granulose, centre-line keeled, with a single row glochids, glochids erect; marginal glochids in 2 rows, erect, two nutlets with the inner glochids 0.5–1 mm long, outer glochids 0.2–0.5 mm long (Fig. 3N); two other nutlets with short glochids, inner glochids less than 0.5 mm long, outer glochids reduced to 0.1–0.2 mm or tuberculate (Fig. 3M); nutlets thin, ca. 0.5 mm thickness (Fig. 3O); abaxially granulose, cicatrix narrow lanceolate, ca. 1 mm long, located in the base of nutlets, adaxial keel ca. 1.5 mm long (Fig. 3P).

Distribution and ecology

The new species is currently known only from its type locality in Dahongliuxia Village, Balikun County, Xinjiang Province, China. It grows in gravel desert at an elevation of 840 m above sea level.

Phenology

Flowering and fruiting from May to July.

Etymology

The specific epithet refers to the appearance of new species, stems frequently branched at middle and above and nearly horizontal spreading.

Vernacular name

Simplified Chinese: 展枝鹤虱 (Chinese pinyin: zhǎn zhī hè shī).

Conservation status

Based on the current survey data, we have only found a single population of the new species at its type locality, Dahongliuxia Village, Balikun County, Xinjiang Province, China. Data for the Lappula effusa were still insufficient to assess its conservation status. According to the IUCN Criteria (IUCN 2022), the conservation status of this new species is temporarily assessed as Data Deficient (DD) until more information becomes available.

Notes

Based on the classification of Lappula by Ovczinnikova (2005), the new species L. effusa should belong to the sect. Microcarpae (M. Pop.) Ovczinnikova, ser. Tianschanicae M. Pop. ex Ovczinnikova, which is characterised by the narrowly subulate gynobase, style surpassing the nutlets by ca. 0.5–1 mm, heteromorphic nutlets with two rows of marginal glochids, disc with centre line keel. Amongst this series, there are approximately seven species (Lappula aktaviensis Popov & Zakirov; L. himalayensis; L. pratensis Ching J.Wang; L. sericata Popov; L. subcaespitosa M Popov ex Golosk.; L. tadshikorum; L. tianschanica Popov & Zakirov). L. effusa most resembles L. himalayensis and L. tadshikorum, sharing similar corolla and gynobase morphology. However, nutlets are always important for identification and classification of Lappula (Popov 1953; Riedl 1967; Zhu et al. 1995; Ovczinnikova 2005) and the new species exhibits distinctive nutlet morphology from the compared species: i.e. relatively compressed fruit, acute ovoid nutlets, narrowly ovate disc and short marginal glochids. Additionally, L. himalayensis and L. tadshikorum are distributed in the mountain areas of Xizang and western Xinjiang (Himalaya and Pamir, usually 1800–4000 m a.s.l.), while L. effusa occurs in the gravel desert of eastern Xinjiang (lower than 1000 m a.s.l.). The detailed differences amongst these three species are provided in Table 1.

Table 1.

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Morphological comparisons of L. effusa L. himalayensis and L. tadshikorum.

Characters	L. effusa	L. himalayensis	L. tadshikorum
Habit	12–28 cm	7–15 cm	30–50 cm
Stem	stem single, erect, branched from middle and above, with spreading hair	stems 4–6, cespitose, ascending or erect, branched from above, with appressed hair	stems 2–3, erect, branched from above, with (semi-) appressed hair
Leave indumentum	abaxial densely spreading hispid, adaxially sparsely hispid or glabrous	abaxial densely appressed pubescent, adaxially sparsely pubescent	abaxial densely spreading hair, adaxially sparsely
Inflorescences	5–10 cm in fruit	3–7 cm in fruit	3–7 cm in fruit
Pedicel	2–3 mm long in fruit	1.5–2 mm long in fruit	1–3 mm long in fruit
Calyx	ca. 2 mm and shorter than the nutlets	ca. 2 mm and shorter than the nutlets	ca. 2 mm and shorter than the nutlets
Corolla	blue or white	blue	blue
Style	surpassing the fruit ca. 0.5 mm and surpassing glochids	surpassing the fruit ca. 0.5 mm and surpassing glochids	surpassing the fruit, but not surpassing glochids
Nutlets	heteromorphic; acute ovoid	heteromorphic; ovoid	heteromorphic; ovoid
Marginal glochids	two rows; 0.5–1 mm long inner glochids erect	two rows; 1.5 mm long; inner glochids often curved	two rows; 1–1.2 mm long; inner glochids curved
Disc of nutlets	narrowly ovate-triangular	ovoid	ovoid
Flowering and fruiting	May to July	June to August	June to July
Elevation	840 m	3700–4200 m	1800–3000 m

Acknowledgements

We would like to thank Dr. Hafiz Muhammad Wariss for modifying the English and two anonymous reviewers for comments on earlier versions of the manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was financially supported by the Natural Science Foundation of Xinjiang Uygur Autonomous Region, China (grant no. 2023D01B02), National Natural Science Foundation of China (grant no. 32300183), the Third Xinjiang Scientific Expedition Program (grant no. 2022xjkk1505) and Shanghai Cooperation Organization Partnership and International Technology Cooperation Plan of Science and Technology Projects (grant no. 2023E01018).

Author contributions

Investigation: DHL, XYZ, SJS, JJW. Data curation: DHL, XYZ, SJS, JJW. Formal analysis: DHL, XYZ. Writing-original draft: DHL. Writing-review and editing: DHL, WJL.

Author ORCIDs

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

Yi-Xin Zhou https://orcid.org/0000-0002-0054-0301

Shu-Jing Shang https://orcid.org/0009-0006-8092-7616

Jia-Ju Wu https://orcid.org/0009-0009-0640-0754

Wen-Jun Li https://orcid.org/0000-0002-2932-0783

Data availability

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

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