Research Article |
Corresponding author: Li Ma ( maliwasps@aliyun.com ) Corresponding author: Qiang Li ( liqiangkm@126.com ) Academic editor: Michael Ohl
© 2024 Jinghong Li, Li Ma, Qiang Li.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Li J, Ma L, Li Q (2024) Five new species of Passaloecus Shuckard (Hymenoptera, Crabronidae) from China, with a key to Chinese species. Journal of Hymenoptera Research 97: 1141-1161. https://doi.org/10.3897/jhr.97.135489
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Based on the key diagnostic characteristics such as petiole, clypeus, and hypersternaulus, five new species of Passaloecus from China are identified: P. clypearcuatus Li & Ma, sp. nov., P. clypeconvexus Li & Ma, sp. nov. P. edentutus Li & Ma, sp. nov., P. margdentatus Li & Ma, sp. nov., and P. sternoleios Li & Ma, sp. nov. Detailed descriptions, diagnoses, and photographs of diagnostic characteristics are provided for these new species. Additionally, a key to the known species of Passaloecus in China is updated and illustrated, further enhancing the understanding of biodiversity within this genus.
Digger wasps, identification key, morphology, Pemphredoninae, taxonomy
In 1837, Shuckard placed Pemphredon insignis within the genus Passaloecus, establishing Passaloecus as a new taxon and designating Passaloecus insignis (Vander Linden, 1829) as its type species (
Passaloecus is represented by 46 species and five subspecies worldwide, with the majority of species distributed in the Palearctic region (30 species and three subspecies) and the Nearctic region (19 species and one subspecies). The Oriental region has a relatively lower diversity, with 13 species and one subspecies, while only one species occurs in the Neotropical region (
This study provides detailed descriptions and illustrations of five new species from China: P. clypearcuatus sp. nov., P. clypeconvexus sp. nov., P. edentutus sp. nov., P. margdentatus sp. nov., and P. sternoleios sp. nov. Additionally, an illustrated key to all known Passaloecus species in China is provided.
The specimens examined in this study were collected using malaise traps, yellow plates, sweep nets, and flight intercept traps, and were deposited in the Insect Collections of Yunnan Agricultural University, Kunming, Yunnan, China (YNAU). Observations were conducted under an Olympus stereomicroscope (SZ Series) with an ocular micrometer. Measurements were taken at 5× magnification, except for body length, which was measured at 2× magnification and subsequently converted. Photographs were captured using a VHX-5000 and edited with Adobe Photoshop 8.0. Morphological terminology follows
AOD Distance from inner eye margin to antennal socket, frontal view;
BL Body length;
EDL Distance between inner eye margins at base of clypeus, frontal view;
EDU Distance between inner eye margins at base of vertex, dorsally;
EL Eye length in lateral view, maximum;
EW Eye width in lateral view, maximum;
EWd Eye width in frontal view, maximum;
HLD Head length in dorsal view, the distance from occipital margin to frons, medially;
HLF Head length in front view, the distance from the clypeal margin to the vertex, medially;
HW Head width, dorsally;
IAD Distance between antennal sockets, frontal view;
LFI Length of flagellomere I;
LFII Length of flagellomere II;
LTI Length of gastral tergum I, maximum, dorsally;
OCD Ocello-occipital distance, distance between posterior margin of hind ocellus and occipital margin, dorsally;
OOD Ocellocular distance, distance between outer margin of hind ocellus and nearest inner orbit, dorsally;
PL Pedicel length;
PLL Petiole length laterally, maximum;
POD Postocellar distance, distance between inner margins of hind ocelli, dorsally;
PW Petiole width, maximum, dorsally;
SL Scape length;
TW Gena width in lateral view, maximum;
WAS Width of antennal socket, frontal view;
WFI Width of flagellomere I;
WFII Width of flagellomere II;
WTI Width of gastral tergum I, maximum, dorsally.
Pemphredon insignis Vander Linden, 1829.
Females of P. multituberculatus Ma & Li, P. petiolatus Ma & Li and P. tuberculiformis Bashir & Ma and males of P. frontirugatus Bashir & Ma, P. labrinigratus Ma & Li, P. monilicornis taiwanus Tsuneki, P. clypearcuatus sp. nov., P. clypeconvexus sp. nov., P. edentutus sp. nov., P. margdentatus sp. nov., and P. sternoleios sp. nov. remain unknown.
1 | Clypeus deeply concave, free margin not produced (Fig. |
P. clypearcuatus Li & Ma, sp. nov. |
– | Clypeus flat or raised, free margin slightly or broadly produced (Fig. |
2 |
2 | Mandible tridentate apically | 3 |
– | Mandible bidentate apically (Fig. |
5 |
3 | Interantennal tubercle narrow and short; free margin of clypeus toothless, females truncate, males nearly arcuate | P. profundesulcatus Bashir & Ma |
– | Interantennal tubercle robust, forming columned or conical projection; free margin of clypeus with three small teeth | 4 |
4 | Pronotal collar without anterior, transverse carina and antero-lateral corner; posterior area of mesopleuron smooth, without rugae; hypersternaulus shallow, short, inconspicuously crenate; gaster distinctly constricted between terga I and II | P. columnaris Ma & Li |
– | Pronotal collar with robust, anterior, transverse carina, antero-lateral corner slightly produced; posterior area of mesopleuron with dense, short, longitudinal rugae; hypersternaulus deep, normal length, distinctly crenate; gaster slightly constricted between terga I and II | P. corniger Shuckard |
5 | Petiole slightly to distinctly longer than wide (Fig. |
6 |
– | Petiole not longer than wide (Fig. |
11 |
6 | Hypersternaulus smooth, not crenate (Fig. |
7 |
– | Hypersternaulus distinctly crenate (Fig. |
8 |
7 | Free margin of clypeus slightly produced, with three distinct, conical teeth medially (Fig. |
P. margdentatus Li & Ma, sp. nov. |
– | Free margin of clypeus broadly produced, with distinct emargination medially (Fig. |
P. edentutus Li & Ma, sp. nov. |
8 | Inner orbital furrow broad, smooth, and with conspicuous, inner marginal carina (Fig. |
P. sternoleios Li & Ma, sp. nov. |
– | Inner and outer orbital furrows lacking; occipital carina narrow, not crenate; notaulus slightly or distinctly impressed; sternum I with several, short, longitudinal rugae | 9 |
9 | Lateral surface of propodeum with dense, slender, oblique rugae anteriorly; sternum I in anterior half with slender, median, longitudinal carina; gaster not constricted between terga I and II | P. birugatus Bashir & Chen |
– | Lateral surface of propodeum smooth, without rugae anteriorly; sternum I without median, longitudinal carina; gaster slightly or distinctly constricted between terga I and II | 10 |
10 | Admedian line slightly impressed; gaster distinctly constricted between terga I and II; gena with dense, fine punctures ventrally; clypeus with dense, midsize punctures | P. petiolatus Ma & Li |
– | Admedian line distinctly impressed; gaster slightly constricted between terga I and II; ventral gena with sparse, fine punctures; clypeus with sparse, fine punctures | P. tuberangustus Bashir & Ma |
11 | Scrobal suture distinct, weakly to distinctly crenate (Fig. |
12 |
– | Scrobal suture absent or weakly impressed, not crenate | 16 |
12 | Omaulus present; ocellar triangle flat | P. turionum Dahlbom |
– | Omaulus absent (Fig. |
13 |
13 | Antero-lateral corner markedly produced; hypersternaulus and episternal sulcus distinctly broadened; frontal median carina absent | 14 |
– | Antero-lateral corner absent (Fig. |
15 |
14 | Females with scutal patches; interantennal tubercle reduced to one point; labrum slightly constricted subapically; pronotal lobe black | P. labrinigratus Ma & Li |
– | Females without scutal patches; interantennal tubercle distinct, slightly elongated; labrum distinctly constricted subapically; pronotal lobe yellow | P. frontirugatus Bashir & Ma |
15 | Occipital carina distinctly crenate; labrum distinctly constricted subapically; anterior part of scutum normal; scutum with sparse, short, longitudinal rugae posteriorly | P. bisulcatus Bashir & Ma |
– | Occipital carina not crenate (Fig. |
P. clypeconvexus Li & Ma, sp. nov. |
16 | Anterior part of scutum normal | 17 |
– | Anterior part of scutum nearly right angle, significantly higher than pronotum | 18 |
17 | Clypeus distinctly convex medially; lateral surface of propodeum with dense, slender, oblique rugae anteriorly; antero-lateral corner moderately to distinctly produced; pronotal lobe white; in females, head from above with temples distinctly convergent posteriorly | P. clypealis Faester |
– | Clypeus nearly flat medially; lateral surface of propodeum smooth anteriorly; antero-lateral corner absent; pronotal lobe brown to black; in females, head from above with temples slightly convergent posteriorly | P. singularis Dahlbom |
18 | Females with scutal patches; in males, posterior area of tergum VI without spinose tubercles | P. turanicus Gussakovskij |
– | Females without scutal patches; in males, posterior area of tergum VI with conspicuous spinose tubercles | 19 |
19 | Propodeal enclosure and posterior surface of propodeum irregularly reticulated | 20 |
– | Propodeal enclosure with irregular rugae or reticulation medially, and with sparse or dense, oblique rugae on each side; posterior surface of propodeum with irregular rugae | 21 |
20 | Pronotal collar with robust, anterior, transverse carina, antero-lateral corner markedly produced; occipital carina not crenate; lateral surface of propodeum with dense, slender, irregular, oblique rugae anteriorly, and with sparse, sturdy, oblique rugae posteriorly; body predominantly black | P. multituberculatus Ma & Li |
– | Pronotal collar without anterior, transverse carina or antero-lateral corner; occipital carina distinctly crenate ventrally; lateral surface of propodeum shiny, nearly smooth anteriorly, and with sturdy reticulation posteriorly; body predominantly yellowish-brown to reddish-brown | P. tuberculiformis Bashir & Ma |
21 | Occipital carina distinctly crenate; parapsidal line weakly impressed; metanotum with sparse, tiny punctures; flagellum mostly yellow | P. koreanus Tsuneki |
– | Occipital carina not crenate; parapsidal line distinctly impressed; metanotum with dense, fine punctures; flagellum dark brown to black | 22 |
22 | Antero-lateral corner markedly produced; propodeal enclosure reticulated medially; frontal median carina weakly impressed; in male, flagellum normal | P. insignis (Vander Linden) |
– | Antero-lateral corner slightly produced; propodeal enclosure irregularly rugose medially; frontal median carina absent; in male, flagellum slightly moniliform | 23 |
23 | Pronotal lobe ivory to yellowish | P. monilicornis monilicornis Dahlbom |
– | Pronotal lobe black | P. monilicornis taiwanus Tsuneki |
Holotype : China • ♀; Yunnan, Shangri-La City, Pudacuo National Park; 27°55'13"N, 99°52'46"E; 3515 m elev.; 16.VII.2022; coll. Zhizhi Liu; sweep net (YNAU). Paratype: China • 1♀; Shaanxi, Baoji City, Tiantai Mountain Scenic Area; 34°17'10"N, 107°10'55"E; 852 m elev.; 8–10.VI.1998; NO.983654; coll. Yun Ma; sweep net (YNAU).
This species differs from the similar species P. labrinigratus Ma & Li, 2012 and other congeners in the following characteristics: free margin of clypeus with two robust, triangular teeth medially, distinctly reflected, area between two teeth deeply concave; scutal patches absent; scutum with sparse, irregular, short, longitudinal rugae posteriorly; scapal hollow coriaceous, shiny and clearly defined; gaster not constricted between terga I and II. Passaloecus labrinigratus has the following characters: free margin of clypeus broadly produced, truncate medially; scutal patches ovate; scutum smooth, without rugae posteriorly; scapal hollow coarsely coriaceous, moderately matte, not clearly defined; gaster slightly constricted between terga I and II.
Female. Measurements. ♀, BL: 7.7 mm; HLF: HW: HLD = 63: 80: 51; HW: EWd: EW: TW: EL = 80: 15: 20: 28: 49; POD: OOD: OCD = 13: 13: 23; SL: PL: LFI: LFII: WFI: WFII = 29: 7: 12: 11: 6: 6; WAS: AOD: IAD = 6: 10: 14; EDU: EDL = 47: 49; PLL: PW: LTI: WTI = 10: 13: 55: 48.
Color pattern. Body black; mandible inner margin pale yellow, remainder reddish-brown to dark brown; labrum ivory medially, central part dark brown; ventral scape ivory to pale yellow; tegula, palpi and forewing veins dark brown; fore tibia and tarsus dark brown; mid leg: inner margin of femur, tibia, and tarsus pale yellow to yellowish-brown, remainder of femur and trochanter dark brown; basal 1/4 of hind tibia dark brown; clypeus with scattered, silvery, short setae.
Head. Mandible bidentate apically, inner tooth large (Fig.
Mesosoma. Pronotal collar with robust, anterior, transverse carina, antero-lateral corner slightly produced (Fig.
Metasoma. Petiole short, not longer than width; dorsal surface of petiole with median, longitudinal groove (Fig.
Male. Unknown.
China (Yunnan, Shaanxi).
The name clypearcuatus is derived from the Latin clype- (= clypeus) and the Latin word arcuatus (= arcuate), referring to the area between two teeth on free margin of clypeus with an arcuate, deep concavity.
Holotype : China • ♀; Xinjiang, Gongliu County, Qiaxi Forest Park; 43°5'44"N, 82°39'18"E; 1713.8 m elev.; 7.VII.2023; coll. Lili Dong; sweep net (YNAU). Paratype: China • 1♀; Xinjiang, Gongliu County, Hetaogou Scenic Area; 43°22'59"N, 82°16'10"E; 913 m elev.; 30.VI.2016; coll. Yicheng Li; sweep net (YNAU).
This species can be distinguished from the similar species P. bisulcatus Bashir & Ma, 2019 and other congeners by: occipital carina not crenate; labrum not constricted subapically; anterior part of scutum nearly right angle, significantly higher than pronotum; scutum smooth, without rugae posteriorly; propodeal enclosure with irregular rugae medially, and with sparse, oblique rugae on each side. Passaloecus bisulcatus has the following characters: occipital carina distinctly crenate; labrum distinctly constricted subapically; anterior part of scutum normal; scutum with sparse, short, longitudinal rugae posteriorly; propodeal enclosure with sturdy, irregular reticulation.
Female. Measurements. ♀, BL: 5.3 mm; HLF: HW: HLD = 53: 54: 36; HW: EWd: EW: TW: EL = 54: 13: 15: 25: 40; POD: OOD: OCD = 8: 9: 12; SL: PL: LFI: LFII: WFI: WFII = 20: 6: 6: 6: 3: 3; WAS: AOD: IAD = 4: 5: 6; EDU: EDL = 32: 22; PLL: PW: LTI: WTI = 6: 10: 38: 36.
Color pattern. Body black; mandible predominantly reddish-brown, basal inner margin grayish-white; palpi and tegula brown to dark brown; scape ventrally and pronotal lobe ivory; fore and mid tibiae and tarsis brown to dark brown; hind leg: tibia brown, except basal 1/4 ivory, tarsus brown; sides of lower frons and clypeus with scattered, silvery, short setae; outer margin of mandible with several golden, long setae.
Head. Mandible bidentate apically, inner tooth large (Fig.
Mesosoma. Pronotal collar with robust, anterior, transverse carina, antero-lateral corner absent (Fig.
Metasoma. Petiole short, not longer than width (Fig.
Male. Unknown.
China (Xinjiang).
The name clypeconvexus is derived from the Latin clype- (= clypeus) and the Latin word convexus (= convex), referring to the clypeus conspicuously convex medially.
Holotype : China • ♀; Tibet, Shigatse City, Yadong County, Shang Yadong Country, Galingang Village; 27°30'28"N, 88°57'40"E; 3445 m elev.; 24.VII.2018; NO.202006141; coll. Shijie Du; sweep net (YNAU). Paratype: China • 1♀; Tibet, Shigatse City, Gyirong County, Gyirong Town, Madun Bridge; 28°23'35"N, 85°19'45"E; 2795 m elev.; 1.VIII.2018; coll. Shijie Du; sweep net (YNAU).
This species differs from P. tuberangustus Bashir & Ma, 2019 and other congeners by these characteristics: hypersternaulus smooth, not crenate; pronotal collar with weak, anterior, transverse carina laterally; lateral surface of propodeum with dense, slender, oblique rugae anteriorly; propodeal enclosure with irregular rugae medially, and sparse, irregular, oblique rugae laterally; mesopleuron coriaceous, posterior area with sparse, indistinct, very short, longitudinal rugae. Passaloecus tuberangustus has the following characters: hypersternaulus conspicuously crenate; pronotal collar without anterior, transverse carina; lateral surface of propodeum smooth and shiny anteriorly; propodeal enclosure with dense, slender, longitudinal rugae, and interspersed with several, short, transverse rugae medially; mesopleuron coarsely coriaceous, posterior area smooth, without rugae.
Female. Measurements. ♀, BL: 6.9 mm; HLF: HW: HLD = 58: 67: 40; HW: EWd: EW: TW: EL = 67: 16: 16: 35: 43; POD: OOD: OCD = 7: 11: 18; SL: PL: LFI: LFII: WFI: WFII = 25: 7: 8: 8: 4: 5; WAS: AOD: IAD = 7: 10: 9; EDU: EDL = 49: 43; PLL: PW: LTI: WTI = 30: 7: 42: 35.
Color pattern. Body black; mandible ivory except apex reddish-brown; palpi ivory to brown; ventral scape and pronotal lobe ivory; tegula and forewing veins dark brown; all tibiae and tarsis brown to dark brown; sides of lower frons and clypeus with scattered, silvery, short setae; outer margin of the mandible with scattered golden, long setae.
Head. Mandible bidentate apically, inner tooth large (Fig.
Mesosoma. Pronotal collar with weak, anterior, transverse carina laterally, without antero-lateral corner (Fig.
Metasoma. Petiole long, conspicuously longer than width; dorsal surface of petiole with indistinct, median, longitudinal groove (Fig.
Male. Unknown.
China (Tibet).
The name edentutus is derived from the prefix e- (= without) and the Latin word dentutus (= dentate), referring to the hypersternaulus smooth, not crenate.
Holotype : China • ♀; Yunnan, Shangri-La City, Jiantang Town, Lindu Village; 27°47'24"N, 99°48'36"E; 3321 m elev.; 13.VII.2022; coll. Huifen Jiang; Yellow Plates (YNAU). Paratype: China • 1♀; Yunnan, Shangri-La City, Jiantang Town, East Ring Road; 27°50'24"N, 99°46'48"E; 3373 m elev.; 14.VII.2022; coll. Lili Dong; sweep net (YNAU).
This species is distinguished from P. tuberangustus Bashir & Ma, 2019 and other congeners by the following characteristics: in female, free margin of clypeus with 3 distinct teeth; hypersternaulus smooth, not crenate; anterior margin of pronotal collar with slender, transverse carina; propodeal enclosure with irregular rugae medially, and sparse, irregular, oblique rugae laterally; pronotal lobe black. Passaloecus tuberangustus has the following characters: in female, free margin of clypeus with shallow emargination medially, in male, free margin of clypeus truncate; hypersternaulus distinctly crenate; pronotal collar without anterior, transverse carina; propodeal enclosure with dense, slender, longitudinal rugae, and interspersed with several, short, transverse rugae medially; pronotal lobe ivory.
Female. Measurements. ♀, BL: 8.5 mm; HLF: HW: HLD = 77: 85: 42; HW: EWd: EW: TW: EL = 85: 20: 25: 50: 58; POD: OOD: OCD = 9: 12: 26; SL: PL: LFI: LFII: WFI: WFII = 35: 7: 11: 10: 6: 7; WAS: AOD: IAD = 6: 7: 7; EDU: EDL = 38: 37; PLL: PW: LTI: WTI = 20: 13: 52: 55.
Color pattern. Body black; mandible ivory except apex reddish-brown; margin of labrum ivory, central part brown; palpi brown to reddish-brown; scape ventrally ivory; tegula and forewing veins dark brown; fore leg: femur reddish-brown basally, tibia and tarsus yellowish-brown to dark brown; mid and hind tibiae and tarsis dark brown; sides of lower frons and clypeus with scattered, silvery, short setae; outer margin of mandible with scattered, golden, long setae.
Head. Mandible with two blunt teeth apically, inner tooth large (Fig.
Mesosoma. Anterior margin of pronotal collar with weak, transverse carina, antero-lateral corner absent (Fig.
Metasoma. Petiole long, conspicuously longer than width (Fig.
Male. Unknown.
China (Yunnan).
The name margidentatus is derived from the Latin marg- (= margin), and the Latin word dentatus (= dental), referring to the free margin of clypeus with three distinct teeth medially.
Holotype : China • ♀; Guangdong, Qingyuan City, Fogang County, Guanyin Mountain; 23°58'12"N, 113°33'49"E; 353 m elev.; 15–16.IX.2007; Yellow Plates (YNAU). Paratype: China • 1♀; Guangdong, Shaoguan City, Nanling National Nature Reserve; 24°54'49"N, 113°2'30"E; 845 m elev.; 23.V.2020; coll. Fei Ye; flight intercept traps (YNAU).
This species differs from P. insignis (Vander Linden, 1829) and other congeners by these characteristics: petiole conspicuously longer than width; inner and outer orbital furrows broad, marginal carina conspicuous, outer orbital furrow distinctly crenate; occipital carina broad, conspicuous crenate; notaulus lacking; sternum I smooth, without carina and rugae. Passaloecus insignis has the following characters: petiole not longer than width; inner and outer orbital furrows lacking; occipital narrow, not crenate; notaulus shallowly impressed, crenate; sternum I with sturdy, median, longitudinal carina and several, short, longitudinal rugae.
Female. Measurements. ♀, BL: 7.8 mm; HLF: HW: HLD = 73: 78: 30; HW: EWd: EW: TW: EL = 78: 20: 26: 33: 60; POD: OOD: OCD = 10: 13: 16; SL: PL: LFI: LFII: WFI: WFII = 28: 6: 7: 7: 5: 5; WAS: AOD: IAD = 6: 5: 7; EDU: EDL = 42: 32; PLL: PW: LTI: WTI = 25: 10: 40: 52.
Color pattern. Body black; mandible ivory except apex yellowish-brown to reddish-brown; central part of labrum with ivory, V-shaped stripe, remainder yellowish-brown; palpi ivory to pale yellow; ventral scape and pronotal lobe ivory; dorsal scape, pedicel, flagellum, tegula, forewing veins yellowish-brown to dark brown; fore leg: apex of trochanter ivory, remainder dark brown, femur dark brown except base and apex yellowish-brown to brown, tibia and tarsus pale yellow to yellowish-brown; mid and hind legs: apex of trochanter, base of femur, ventral tibia and tarsus yellowish-brown, remainder dark brown; sides of lower frons and clypeus with scattered, silvery, short setae; clypeus with sparse, golden, short setae, and interspersed with several golden, long setae apically; labrum with several, golden, long setae apically.
Head. Mandible bidentate apically, inner tooth large (Fig.
Passaloecus sternoleios Li & Ma, sp. nov. (female) A head, frontal view B head, dorsal view C collar, scutum, scutellum and metanotum, dorsal view D thorax, lateral view E propodeum, dorsal view F petiole and tergum I dorsal view G petiole and sternum I ventral view H habitus, lateral view. Scale bars: 1 mm.
Mesosoma. Anterior margin of pronotal collar with strong, arcuate, transverse carina, antero-lateral corner markedly produced (Fig.
Metasoma. Petiole long, conspicuously longer than width, smooth, without carina or rugae (Fig.
Male. Unknown.
China (Guangdong).
The name sternoleios is derived from the Greek stern- (= sternum), and the Greek word leios (= smooth), referring to the sternum I smooth and shiny, without carina or rugae.
The genus Passaloecus comprises 46 species and 5 subspecies globally, primarily distributed across the Palearctic, Nearctic, and Oriental regions. China boasts the highest diversity of this genus, with 18 species and 1 subspecies recorded across its Provinces (
Notably, all 13 known species and one subspecies of Passaloecus in the Oriental region are exclusively distributed in China, with no records found in other Oriental countries, such as those in Southeast Asia (
From a global perspective, further collection and research efforts are essential. Although only one species of the Passaloecus genus is currently recorded in the Neotropical region, the climatic conditions there are highly conducive to species diversification, suggesting the potential existence of additional undiscovered species (
We extend our heartfelt thanks to Wojciech J. Pulawski (California Academy of Sciences, California) for providing us with numerous valuable references, and to Prof. Zhiqiang Li and Dr. Fei Ye from the Institute of Zoology, Guangdong Academy of Sciences, for supplying the specimens. We also express our sincere gratitude to the section editor and all anonymous reviewers for their valuable and constructive comments, which have greatly improved this article. This work was supported by the National Natural Science Foundation of China under Grant number 32270485 and the Agricultural Basic Research joint project of Yunnan Province under Grant number 202101BD070001-004.