Research Article
Research Article
A new species of Pima Hulst, 1888 from China (Lepidoptera, Pyralidae, Phycitinae), with a key to Holarctic species
expand article infoLinlin Yang, Yingdang Ren
‡ Henan Academy of Agricultural Science, Zhengzhou, China
Open Access


Pima tristriata sp. nov. is described as new to science based on specimens collected from the Ningxia Hui Autonomous Region, China, and P. boisduvaliella (Guenée, 1845) is also treated here for comparison. DNA barcodes of the two species are provided, together with a neighbor-joining tree for species delimitation. A key to the Holarctic species and a distribution map of the Chinese species are presented.


COI, key, new species, Pima boisduvaliella, Pima tristriata, snout moths, taxonomy


The genus Pima was established by Hulst (1888) with Pima fosterella Hulst as the type species. Ragonot (1889, 1893) referred P. fosterella and the other congeneric American species to Epischnia Hübner, 1825. Heinrich (1956) revised the genus Pima from America, pointing out that Epischnia as defined by Ragonot was a composite of several disparate species and that none of them agreed with the type species of Epischnia, and transferred eight species to Pima. Neunzig (2003) treated nine species of Pima from North America, described one new species and proposed two synonyms. Leraut (2014) treated five species from Europe, including one new species and two new combinations. Tsvetkov (2016) described P. transfusor Tsvetkov from the South Urals. Moreno and Gastón (2017) transferred four species to Pima. Falck et al. (2019) described P. tricolorella Falck, Karsholt & Slamka from the Canary Islands of Spain. Slamka (2019) reviewed the genus in Europe, synonymized Palloria Amsel with Pima and Pima leucomixtella (Ragonot) with Pima christophori (Ragonot), transferred Epiepischnia keredjella Amsel and Epischnia trifidella Zerny to Pima, and described three new species. Twenty-four species have hitherto been assigned to Pima worldwide, mainly distributed in North America and Europe.

Two species, Pima boisduvaliella (Guenée) and P. trifidella (Zerny) were reported from China before this study. We herein describe one new species, Pima tristriata sp. nov., provide DNA barcodes of the new species and P. boisduvaliella (Guenée), and a neighbor-joining tree covering seven species for species delimitation. A key to the known Holarctic species of the genus Pima is also provided.

Materials and methods

The examined specimens in this study were collected by light traps in the Ningxia Hui Autonomous Region, China. Morphological terminology follows Heinrich (1956). Genitalia and wings were dissected and mounted according to the methods introduced by Li (2002). Illustrations were prepared using a Leica DM750 microscope, and refined in Photoshop CS4 software. Photographs of adults were taken with a Leica M205A stereo microscope. The cartographic illustration was made using DIVA-GIS 7.5 (Hijmans et al. 2005). All specimens examined, including the holotype of the new species, are deposited in the Insect Collection of Nankai University, Tianjin, China (NKU).

DNA was extracted from dry adult specimens using Qiagen DNeasy Blood & Tissue Kit, with the genitalia mounted on slides as vouchers. Samples were amplified using the primers LCO1490 and HCO2198 (Folmer et al. 1994) in 25 μl reaction volume: 0.75 μl of each primer (10 mM; Sangon Biotech), 2 μl DNA template, 12.5 μl mixture (KOD One PCR Master Mix; TOYOBO), and 9 μl ddH2O. PCR reaction conditions used were as follows: 35 cycles of 98 °C for 10 s, 55 °C for 5 s, 68 °C for 1 s; then a 4 °C hold. A weak electrophoretic band of the new species was obtained, and the PCR product was recovered (SanPrep Column DNA Gel Extraction Kit; Sangon Biotech) and cloned (Hieff CloneTM Zero TOPO-TA Cloning Kit; Sangon Biotech). Positive plasmids were sent to Sangon Biotech (Shanghai, China) for sequencing.

Genetic distance estimation and neighbor-joining analysis were conducted in MEGA X using the Kimura 2-Parameter model. Thirty-eight sequences were used in the analyses: one new sequence from a paratype of P. tristriata sp. nov. (GenBank accession number MT749678) and three new ones from Chinese specimens of P. boisduvaliella (GenBank accession numbers MT734539, MT734540, MT734541), the others from BOLD (Ratnasingham and Hebert 2007). The extreme values of the interspecific and intraspecific distances were presented in Table 2, and the NJ tree was shown in Fig. 8.


Pima Hulst, 1888

Pima Hulst, 1888: 114. Type species: Pima fosterella Hulst, 1888, by original designation and monotypy

Palloria Amsel, 1961: 362. Type species: Palloria bicornutella Amsel, 1961

Diagnostic characters. Pima is characterized by the male basal few flagellomeres shallowly incurved and containing a row of minute, tooth-like spines (Figs 1a, 2a), third segment of the labial palpus projected forward (Figs 1b, 2b); the forewing usually having a white subcostal streak (absent in P. keredjella, P. milka, P. parkerella, P. pempeliella, P. transfusor and P. tristriata sp. nov.), with 11 veins (Figs 1c, 2c), R2 approximate to the stalk of R3+4 + R5, R3+4 stalked with R5 of less than half their lengths, M2, M3 and CuA1 free; the hindwing with 10 veins (Figs 1c, 2c), Rs and M1 shortly stalked, M2 and M3 stalked for over half their length, CuA1 and M2+ M3 shortly stalked; apical process of gnathos short and stout, transtilla absent, the broad costa of the narrowed valva with a blunt, slightly forked apex (more pointed and not forked in P. christophori, P. leucoloma, P. pempeliella, and P. trifidella), the uncus with a broad base and a short pair of lateral lobes, the aedeagus with two stout cornuti (one cornutus in P. trifidella) in male genitalia (Figs 3, 4); the ductus bursae ribbon-like, the stout corpus bursae scobinate-granulate and usually with sclerotized patches or folds in female genitalia (Figs 5, 6).

Figures 1, 2. 

Adult Pima species. 1 P. tristriata sp. nov., holotype, male 1a dorsal view of head, holotype, male 1b lateral view of head, holotype, male 1c wing venation, paratype, female, DYL01090 2 P. boisduvaliella, female 2a dorsal view of head, male 2b lateral view of head, male 2c wing venation, female, WYQ13200. Scale bars: 5.0 mm.

Figures 3–6. 

Genitalia of Pima species. 3, 4 Male genitalia 3 P. tristriata sp. nov., paratype, YLL18044 4 P. boisduvaliella, DYL01090 5, 6 Female genitalia 5 P. tristriata sp. nov., paratype, LJY10110 6 P. boisduvaliella, DYL00331. Scale bars: 0.5 mm.

Pima resembles Epischnia Hübner, but they can be separated by the following characters: in Pima, the male flagellum with a row of tooth-like spines near the base, the labial palpus with terminal two segments approximately of equal length; male genitalia with a broad, apically slightly forked costa, and two stout cornuti in the aedeagus; female genitalia with a strongly sclerotized, funnel-shaped antrum, the corpus bursae scobinate-granulate throughout and with sclerotized patches or folds. Whereas, in Epischnia, the male flagellum lacks a tooth-like spine, the third of the labial palpus is less than half the length of the second; the costa is weak and not forked at the apex, and the aedeagus has a bunch of spinules in the male genitalia; the antrum is weak or represented by a band-shaped plate, the corpus bursae is smooth on the inner surface except for one big sclerotized plate or a line of small thorns and one bunch of spinules in the female genitalia.

Key to Holarctic species of the genus Pima

1 Forewing with distinct white subcostal streak 2
Forewing with obscure white subcostal streak or absent 17
2 Forewing ground color creamy-whitish, with a distinct longitudinal brown streak under white subcostal streak (Amsel 1961: pl. 3, fig. 181; Slamka 2019: pl. 22, fig. 152) P. keredjella
Forewing ground color yellowish, grayish or brown, without distinct streak under white subcostal streak 3
3 White subcostal streak conspicuously developed only form base of costa to the antemedial line (Falck, Karsholt and Slamka 2019: figs 5, 6) P. tricolorella
White subcostal streak well developed along whole length of forewing 4
4 Costa of valva has a more pointed and not forked apex 5
Costa of valva has a blunt, slightly forked apex 7
5 Aedoeagus has a single cornutus (Slamka 2019: pl. 78, fig. 156) P. trifidella
Aedoeagus has two cornuti 6
6 Forewing ground color grayish, with faint postmedial line (Vives Moreno and Gastón 2017: fig. 19; Slamka 2019: pl. 22, figs 154a–e) P. leucoloma
Forewing ground color pale yellow, without transverse line (Leraut 2014: pl. 41 fig. 8; Slamka 2019: pl. 22, figs 155a–d) P. christophori
7 Corpus bursae with a slug-shaped sclerotization 8
Corpus bursae without the above sclerotization 13
8 Two cornuti ca equal thickness 9
Shorter cornutus broader than the longer one 10
9 Corpus bursae heart-shaped (Fig. 6) P. boisduvaliella
Corpus bursae oblong (Heinrich 1956: fig. 777) P. albiplagiatella
10 Shorter cornutus significantly shorter than the longer one (Leraut 2014: text fig. 121c; Slamka 2019: pl. 73, fig. 146a, pl. 74, fig. 146b–d); Corpus bursae ca 1.5× length of its medial width (Leraut 2014: fig. 122a; Slamka 2019: pl. 154, fig. 146) P. tabulella
Shorter cornutus slightly shorter than the longer one; Corpus bursae more than double length of its medial width 11
11 Shorter cornutus broad at base, abruptly tapered to apex (Slamka 2019: pl. 74, figs 148) P. marocana
Shorter cornutus gradually tapered to apex 12
12 Gnathos-arms stouter (Slamka 2019: pl. 74, fig. 147); Corpus bursae sclerotized in posterior three-quarters (Leraut 2014: text fig. 122c; Slamka 2019: pl. 155, figs 147a, b) P. aureliae
Gnathos-arms narrower (Slamka 2019: pl. 75, fig. 149); Corpus bursae sclerotized in posterior three-quarters (Slamka 2019: pl. 155, fig. 149a, pl. 156, fig. 149b–c) P. yllai
13 Corpus bursae oblong 14
Corpus bursae nearly rounded 15
14 Corpus bursae without hump-shaped protuberance (Neunzig 2003: text fig. 3) P. occidentalis
Corpus bursae with a sclerotized hump (Slamka 2019: pl. 156, figs 150a, b) P. vilhelmseni
15 Forewing pale (Neunzig 2003: pl. 1, fig. 1); corpus bursae not sclerotized anteriorly (Heinrich 1956: fig. 783; Neunzig 2003: fig. 2c) P. fosterella
Forewing darker; corpus bursae sclerotized anteriorly 16
16 Forewing salmon pink below white subcostal streak (Neunzig 2003: pl. 1, fig. 6); antrum subovate (Heinrich 1956: fig. 780) P. fulvirugella
Forewing dark gray to blackish brown below white subcostal streak (Neunzig 2003: pl. 1, fig. 7; Leraut 2014: pl. 41, fig. 11); antrum funnel-shaped (Heinrich 1956: figs 778, 779) P. albocostalialis
17 Forewing with faint antemedial and postmedial lines 18
Forewing without transverse lines 22
18 Costa of valva not forked at apex 19
Costa of valva slightly forked at apex 20
19 Juxta V-shaped, aedeagus significantly shorter than the valva, clasper present (Roesler 1990: fig. 9); apophyses posteriores slightly shorter than anteriores (Roesler 1990: fig. 10) P. milka
Juxta U-shaped, aedeagus as long as the valva, clasper absent (Roesler 1990: fig. 11; Slamka 2019: pl. 76, fig. 153); apophyses posteriores significantly shorter than anteriores (Roesler 1990: fig. 12; Slamka 2019: pl. 157, fig. 153) P. pempeliella
20 Forewing with longitudinal grayish black streaks along costa and dorsum 21
Forewing without longitudinal fuscous streaks (Tsvetkov 2016: fig. 1) P. transfusor
21 Forewing with a longitudinal grayish black streaks along lower margin of cell (Fig. 1); costa projected beyond apex of valva (Fig. 3); corpus bursae heart-shaped (Fig. 5) P. tristriata sp. nov.
Forewing without longitudinal streaks along lower margin of cell (Neunzig 2003: pl. 1, fig. 10); costa not projected beyond apex of valva (Heinrich 1956: fig. 306); corpus bursae more rounded, with a projecting shield at junction with ductus bursae (Heinrich 1956: fig. 782) P. parkerella
22 Forewing more nearly uniform, without contrasting longitudinal lines (Neunzig 2003: pl. 1, figs 11, 12) P. fergusoni
Forewing more black along veins (Neunzig 2003: pl. 1, figs 8, 9) P. granitella

Pima tristriata sp. nov.

Figures 1, 3, 5

Type material

Holotype : China: • ♂; Shapotou (37°31'N, 105°10'E), Zhongwei, Ningxia Hui Autonomous Region; alt. 1140 m; [?]-v-1985; Guo-Dong Ren leg. Paratypes: China: • 7♂; same data as the holotype; genitalia nos. DYL01079, DYL01080, RYD04466 • 3♂, 2♀; same data as the holotype except dated 23-iv-1987; genitalia nos. YLL18042♂, YLL18044♂ • 2 ♀; Gantang (37°27'N, 105°32'E), Zhongwei, Ningxia; 23-v-1987; Guo-Dong Ren leg.; genitalia no. DYL01090.


The new species can be easily distinguished from its congeners in having one longitudinal grayish black streak along the costa, dorsum, and lower margin of cell respectively, whereas, most of the other congeners have a white subcostal streak. It is superficially similar to P. parkerella (Schaus), but with differences in genitalia: juxta with globular lateral lobes, costa projected beyond apex of valva, and corpus bursae heart-shaped in the new species; juxta with short finger-like lateral lobes, costa terminated at end of valva, and corpus bursae rounded in P. parkerella. It resembles Pima boisduvaliella (Guenée) in genitalia except for some slight differences: lateral lobes the juxta is globular, the vinculum is ca 2× length of its greatest width, the aedeagus is approximately equal to valva in length in the male genitalia, and the corpus bursae has an irregular sclerotized plate in the female genitalia. In P. boisduvaliella, lateral lobes the juxta is slender, finger-like, the vinculum is ca 1.5× length of its greatest width, and the aedeagus is 1.2× length of valva in the male genitalia; the corpus bursae has a couple of tortuous, sclerotized plates in the female genitalia.


Adult (Fig. 1). Wingspan 25.5–31.0 mm. Head (Fig. 1a, b) grayish white. Antenna grayish white, scape ca 1.5× as long as wide, flagellum of male with short cilia, of female pubescent. Labial palpus of male grayish white mixed with a few brown scales, of female brown mixed with a few grayish white scales; first and second segments obliquely upturned, third second projected forward; third segment as long as second, twice as long as first. Maxillary palpus minute, grayish brown, in form of an aigrette. Patagium, tegula and thorax pale yellow, mottled a few brown scales. Forewing yellow, costa dorsum and lower margin of cell overlaid with a longitudinal grayish black streak respectively, more or less peppering of whitish scales; some scattered black dotting along veins and termen; antemedial line white, arched, white, from costal 1/5 slightly oblique to dorsum 1/4, inner bordering ashy black on lower half, outer edging of grayish brown; postmedial line indistinct; discal spots brown, separated; postmedial line black, obscure; cilia yellowish write. Hindwing pale gray, cilia grayish white.

Male genitalia (Fig. 3). Uncus oval, lateral margins enfolded at distal half. Apical process of gnathos conical, ca 1/3 length of uncus. Transtilla absent. Valva narrow, 5× as long as wide; clasper a narrowed triangular process, with a globular, haired base; costa stout, slightly longer than and ca 2/3 width of valva, its apex blunt, slightly forked; sacculus ca 2/5 length of valva, broader at base, tapering toward pointed apex, bearing dense, spine-like hairs along ventral margin. Juxta a broad, semicircular plate, with a pair of short, globular lateral lobes. Vinculum twice as long as its greatest width, narrowly rounded anteriorly. Aedeagus nearly as long as valva, slightly curved towards base, with a tuft of granulations near base; Cornuti two stout thorns, longer one slightly less than half length of aedeagus. Culcita one pair of long hair tufts, 2/3 length of valva.

Female genitalia (Fig. 5). Ovipositor triangular, 3× as long as wide. Apophyses posteriores slender, 3/4 length of apophyses anteriores. Eighth tergite 2/3 length of its width. Antrum strongly sclerotized, funnel-shaped, broader than eighth segment. Ductus bursae sclerotized, 1.2× as long as corpus bursae, of nearly equal width throughout, slightly broader at junction with corpus bursae. Corpus bursae heart-shaped, scobinate-granulate on inner surface, with two sclerotized patches: one oval sclerotized plate near middle; one irregular large plate from junction with ductus bursae to anterior 1/3, its posterior half smooth, forming a shallow fold along its edge, anterior half granulated and wrinkled. Ductus seminalis from posterior margin of corpus bursae.

DNA barcode

One DNA barcode from a female paratype was obtained and deposited in GenBank (accession numbers: MT749678), DNA voucher slide no. DNAYLL18119.


The specific name is derived from the Latin prefix tri-, meaning three, and the Latin word striatus, meaning streak, referring to three grayish black streaks on the forewing.


China (Ningxia).

Host plant


Pima boisduvaliella (Guenée, 1845)

Figures 2, 4, 6

Epischnia boisduvaliella Guenée, 1845: 319.

Anerastia farrella Curtis, 1850: 114.

Myelois lafauryella Constant, 1865: 189.

Pima boisduvaliella (Guenée): Hannemann 1964: 180.


Adults (Fig. 2) with wingspan 15.0–22.0 mm. Pima boisduvaliella is characterized by the yellowish brown forewing with a white subcostal streak; the elongate valva with a well-developed costa that produced and weakly notched apically, the broad semicircular juxta with a pair of short, finger-like lateral lobes, the V-shaped vinculum ca 1.5× length of its greatest width, and the aedeagus with two thorns that slightly less than half the length of the aedeagus in the male genitalia (Fig. 4); the rounded antrum, the heart-shaped corpus bursae with dense microtrichia in anterior 1/3, with a small oval sclerotized plate and a couple of tortuous, sclerotized plates in the female genitalia (Fig. 6).

Three DNA barcodes were obtained and deposited in GenBank: a male collected on August 19, 2007 at alt. 2178 m in Mt. Xinglong, Yuzhong County, Gansu Province, accession no. MT734539, DNA voucher slide no. DNAYLL18043; a male collected on July 24, 2013 at alt. 1461 m in Habahu, Yanchi County, Ningxia Hui Autonomous Region, accession no. MT734540, DNA voucher slide no. DNAYLL18076; a male collected on August 3, 2010 at alt. 1836 m in Shuimogou, Mt. Helan, Alxa Zuoqi, Inner Mongolia Autonomous Region, accession no. MT734541, DNA voucher slide no. DNAYLL18118.


China (Gansu, Hebei, Inner Mongolia, Liaoning, Ningxia, Qinghai, Shaanxi, Shanxi, Xinjiang, Xizang) (Fig. 7), Europe (Slamka 2019: 128, fig. 145), Canada, USA.

Figure 7. 

Geographical distribution of Pima in China: P. tristriata sp. nov. (triangle), P. boisduvaliella (circle); P. trifidella (square).

Host plants

Leguminosae: Anthyllis vulneraria L., Astragalus dasyanthus Pall., Astracantha arnacanthoides, Lathyrus japonicus Willd., Lotus corniculatus L., Ononis spinosa L., O. arvensis L., Hibiscus esculentus (Heinrich 1956; Leraut 2014; Slamka 2019).


Pima is a genus containing 25 species of which 15 are Palaearctic (Roesler 1990; Leraut 2014; Tsvetkov 2016; Vives Moreno and Gastón 2017; Slamka 2019), nine species are Nearctic (Heinrich 1956; Neunzig 2003), and two are Afrotropical (Joannis 1927) (Table 1). Species of Pima might be expected to occur at higher elevations, as most of them were recorded from mountainous areas. In China (Fig. 7), P. boisduvaliella is mainly distributed in the north, but also occurred in the west, such as Xinjiang and Tibet; P. trifidella is distributed in Xinjiang; and P. tristriata sp. nov. is only found in Zhongwei, Ningxia. Adults of the two species were collected from mountain areas with altitudes ranging from 900 m to 3050 m.

Table 1.

Distribution of the Pima species in the worldwide .

Species Distribution
P. albiplagiatella southeastern Canada and northeastern USA
P. albocostalialis southwestern Canada, Pacific Coast states and Rocky Mountain states of USA
P. aureliae Tunisia
P. boisduvaliella from Europe to Central Asia, Southern Canada and Northern USA
P. christophori Armenia, Georgia, Iran, Turkey, Turkmenistan
P. difficilis Mozambique
P. fergusoni Oregon and California of USA
P. flavidorsella Mozambique
P. fosterella western Canada and USA
P. fulvirugella south central and southwestern Canada and Northern California
P. granitella Rocky Mountain and Pacific Coast states of USA
P. keredjella Iran
P. leucoloma Crimea, Croatia, Cyprus, Greece, Italy, Lebanon, Spain, Syria, W Turkey, Tunisia
P. marocana Morocco
P. milka Iran
P. occidentalis Rocky Mountain and Pacific Coast states of USA
P. parkerella Montana of Canada
P. pempeliella Morocco
P. tabulella Altai Republic, NW Mongolia, Turkmenistan
P. transfusor South Urals
P. tricolorella Spain
P. trifidella China
P. tristriata sp. nov. China
P. vilhelmseni Libya, Morocco, Tunisia
P. yallai Morocco, Tunisia

The genetic distance analysis was made based on the pairwise analysis of 38 sequences. According to the NJ bootstrap consensus tree (Fig. 8), ten well-supported clusters of Pima were revealed: P. tristriata sp. nov. is clearly distinguished from its congeners, and this is highly consistent with the morphological analysis; three specimens (LBCG348-08, BBLPD956-10, LCHP302-07) labeled P. albiplagiatella, two specimens (GBMAB2238-15, LBCG1304-09) labeled P. fosterella, and an additional two unidentified specimens (SSKUC2508-15, SSKUC156-15) might represent two unnamed species, as members show higher divergences with P. albiplagiatella and P. fosterella, here treated as P. albiplagiatella sp. inquirenda and P. fosterella sp. inquirenda. Sequence divergences among individuals (Table 2) indicated that minimal interspecific distances range from 1.7 to 2.2%, and the maximal intraspecific distances range from 0 to 1.2%. The present analysis is limited by the relatively small number of species that have been sequenced, and further study is necessary to determine the boundaries of intraspecific and interspecific distances, and whether the minor morphological difference is intraspecific variation or interspecific difference.

Figure 8. 

Neighbor-joining tree deduced from the cytochrome c oxidase subunit I (COI) gene sequences using MEGA X. Sequences were corrected with the Kimura two-parameter substitution model. Codon positions included were 1st + 2nd + 3rd + non-coding. Values represented at the nodes of branches are bootstrap values (1000 replicates).

Table 2.

Percentage of divergence in the cytochrome c oxidase subunit I (COI) gene sequences of the Pima species.

1 2 3 4 5 6 7 8 9 10 11
1 Epischnia illotella
2 E. prodromella 7.4
3 Pima albiplagiatella 12.8–13 9.4–9.9 0–1.2
4 P. albiplagiatella (sp. inquirenda) 12.1–12.5 9.2–9.9 5.4–6.2 0–0.8
5 P. fosterella 11.1–12.5 9.4 1.7–2.2 5.2–6.4 0
6 P. fosterella (sp. inquirenda) 11.4–12.3 9.0–9.2 6.2–7.5 3.5–4.7 5.4–6.0 0.3–1.2
7 P. albocostalialis 12.9 9.4 5.0–5.9 5.2–5.7 5.4–5.6 6.0–6.4
8 Pima sp. 11.3 9.4 5.2–5.9 4.7–5.5 5.4–5.7 5.9–6.0 2.6 0
9 P. boisduvaliella 11.6–12.3 8.4–9.2 2.3–3.2 4.5–5.5 2.0–3.1 5.7–6.1 5.6–6.2 5.4–5.9 0–0.6
10 P. occidentalis 11.7–12.1 8.5–8.7 5.4–6.4 3.9–4.9 5.5–5.8 4.6–5.2 4.7–5.0 5.9 5–5.9 0.2–0.9
11 P. parkerella 14.4 12 8.1–9.1 7.6–7.8 8 7.3–8.2 6.5 7.3 8.1–9.2 6.2–6.4
12 P. tristriata sp. nov. 15.6 13.6 13.3–13.8 15.5–15.9 13.3–13.6 15.9–16.4 15.1 15.1 12.5–13.4 15.3–15.5 18.6


We express our thanks to Dr Guodong Ren (Hebei University, Baoding, Hebei, China) for presenting us with the specimens. We are grateful to Dr Houhun Li (Nankai University, Tianjin, China), Dr František Slamka (Bratislava, Slovakia), and Dr Colin W. Plant (Bishops Stortford, UK) for their constructive suggestions of the manuscript, and Dr Eugene V. Tsvetkov (Saint-Petersburg, Russia) for providing valuable literature. This study was supported by the National Natural Science Foundation of China (31702034 and 31172141) and the Basic Scientific research project of Henan Academy of Agricultural Sciences (2020ZC33 and 2018YQ18).


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