Research Article |
Corresponding author: Cheng-De Li ( lichengde0608@sina.com ) Academic editor: Rikjan Vermeulen
© 2024 Ke-Yi Wang, Fabian A. Boetzl, Jürgen Wiesner, Cheng-De 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:
Wang K-Y, Boetzl FA, Wiesner J, Li C-D (2024) New records and revised distribution of tiger beetles in China (Coleoptera, Cicindelidae). ZooKeys 1210: 29-59. https://doi.org/10.3897/zookeys.1210.127753
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Based on the examination of specimens housed in several scientific collections, we expand the known tiger beetle fauna of China, and eight species are recorded from China for the first time. The occurrence of Cicindela (Cicindela) sachalinensis raddei Morawitz, 1863 in Shanxi Province, and Neocollyris (Neocollyris) saphyrina (Chaudoir, 1850) in China are re-established. We provide distribution maps and habitus photographs of examined specimens for the newly recorded and revised species. We also discuss potential research hotspots for future taxonomic studies of tiger beetles in China.
China, distribution, new records, taxonomy, tiger beetles
Tiger beetles (Coleoptera, Cicindelidae) are a globally distributed family that occupies almost all terrestrial ecosystems (
There has been limited research on the overall distribution of tiger beetles in China (
This article summarizes data from tiger beetle specimens in several universities and research institutes across China, along with those collected by the corresponding author’s research team. Through these specimens, we update and expand the known distribution of some species of tiger beetle throughout China. Some provinces included here have lacked new records of tiger beetles for many years. The results expand our understanding of the tiger beetle fauna of China.
In addition to these specimen data, other records were extracted from
Photographs of the habitus of specimens were taken with a Canon EOS M6 Mark II. A Godox TT350c Flash was used as a light source. Helicon Focus v. 7 was used for image stacking and all images were further processed in Adobe Photoshop CS6.
The specimens reported below are deposited in the following collections:
NEFU Northeast Forestry University, Harbin, China
Distribution maps were generated using ArcGIS v. 10.2. Map data were sourced online from http://xzqh.mca.gov.cn/map (Fig.
1. | Heilongjiang | 2. | Jilin |
3. | Inner Mongolia | 4. | Xinjiang |
5. | Liaoning | 6. | Beijing |
7. | Hebei | 8. | Tianjin |
9. | Gansu | 10. | Shanxi |
11. | Ningxia | 12. | Shandong |
13. | Qinghai | 14. | Shaanxi |
15. | Henan | 16. | Xizang |
17. | Jiangsu | 18. | Anhui |
19. | Shanghai | 20. | Hubei |
21. | Sichuan | 22. | Chongqing |
23. | Zhejiang | 24. | Jiangxi |
25. | Hunan | 26. | Guizhou |
27. | Fujian | 28. | Yunnan |
29. | Taiwan | 30. | Guangxi |
31. | Guangdong | 32. | Hong Kong |
33. | Macao | 34. | Hainan |
Family Cicindelidae Latreille, 1802
Tribe Cicindelini Latreille, 1802
Subtribe Cicindelina Latreille, 1802
Cicindela anchoralis:
Abroscelis anchoralis anchoralis:
Liaoning (
Habitus photographs A Abroscelis anchoralis anchoralis (Chevrolat, 1845) B Apterodela (Apterodela) bivirgulata bivirgulata (Fairmaire, 1889) C Callytron nivicinctum (Chevrolat, 1845) D Calomera chloris (Hope, 1831) E Calomera plumigera scoliographa (Rivalier, 1953) F Cicindela (Cicindela) campestris pontica Fischer, 1828. Scale bars: 5 mm.
Guangdong, Zhanjiang, Min’an Town, 20°57'28"N, 110°15'6"E, 1 m, leg. H.B. Liang and X.L. Huang, 28.vi.2014, 1 male (
Distribution maps. Green indicates records with previously known distribution based on published data, blue indicates new records, red stars indicate the collection sites of the examined specimens A Abroscelis anchoralis anchoralis (Chevrolat, 1845) B Apterodela (Apterodela) bivirgulata bivirgulata (Fairmaire, 1889) C Callytron nivicinctum (Chevrolat, 1845) D Calomera chloris (Hope, 1831) E Calomera plumigera scoliographa (Rivalier, 1953) F Cicindela (Cicindela) campestris pontica Fischer, 1828.
China (Guangdong, Liaoning, Beijing, Hebei, Shandong, ?Jiangsu, Zhejiang, ?Fujian, Hainan, Hong Kong, Macao, Taiwan).
New provincial record for Guangdong.
Subgenus Apterodela Rivalier, 1950
Cicindela bivirgulata:
Cylindera (Apterodela) lobipennis:
Apterodela (Apterodela) bivirgulata:
Shandong (
Jiangsu, leg. unknown, 18.iv.1923, 1 male (
China (Shandong, Henan, Hubei, Anhui, Jiangxi, Jiangsu, Shanghai, Chongqing, Zhejiang, Fujian).
Additional specimens contribute to a better understanding of the distribution and a more robust identification of A. (A.) b. bivirgulata.
Cicindela nivicincta:
Callytron nivicinctum:
Liaoning (
Guangxi, Longzhou, Mount Daqing, leg. J.K. Yang, 15.v.1963, 1 female (
China (Guangxi, Liaoning, Jiangsu, Zhejiang, Shanghai, Fujian, Guangdong, Hainan, Hong Kong, Macao), South Korea, Japan, Cambodia, Vietnam.
New provincial record for Guangxi.
Cicindela chloris:
Xizang, Linzhi, Xiachayu, 28°29'59"N, 97°1'3"E, leg. J. Wu, 28.vii.2014, 1 female (
China (Xizang), Afghanistan, Pakistan, Nepal, Bhutan, India, Laos.
New state record for China and new provincial record for Xizang.
Calomera plumigera scoliographa:
Yunnan (
Guangxi, Jinxiu, Dazhang, leg. unknown, ?.v.1976, 1 male (
China (Guangxi, Yunnan, Hainan), Vietnam, Laos, Cambodia, Malaysia, Thailand.
New provincial record for Guangxi.
Subgenus Cicindela Linnaeus, 1758
Cicindela campestris pontica:
Xinjiang, Altai, Forest Region of Ashan Forest Management Bureau, leg. unknown, v.1981, 1 female (
China (Xinjiang), Ukraine, Bulgaria, Turkey, Georgia, Azerbaijan, Kazakhstan, Russia.
New state record for China and new provincial record for Xinjiang.
Cicindela raddei:
Cicindela sachalinensis:
Cicindela sachalinensis raddei:
Cicindela (Cicindela) sachalinensis raddei:
Heilongjiang (
Habitus photographs A Cicindela (Cicindela) sachalinensis raddei Morawitz, 1863 B Cicindela (Cicindela) transbaicalica hamifasciata Kolbe, 1886 C Cosmodela separata (Fleutiaux, 1893) D Cylindera (Cylindera) obliquefasciata (Adams, 1817) E Cylindera (Eriodera) albopunctata (Chaudoir, 1852) F Cylindera (Eugrapha) contorta contorta (Fischer, 1828). Scale bars: 5 mm.
Shanxi, Mount Wutai, Dongtai, 2400 m, leg. T.S. Li, 13.vi.1964, 1 female (
Distribution maps. Green indicates records with previously known distribution based on published data, blue indicates new records, red stars indicate the collection sites of the examined specimens A Cicindela (Cicindela) sachalinensis raddei Morawitz, 1863 B Cicindela (Cicindela) transbaicalica hamifasciata Kolbe, 1886 C Cosmodela separata (Fleutiaux, 1893) D Cylindera (Cylindera) obliquefasciata (Adams, 1817) E Cylindera (Eriodera) albopunctata (Chaudoir, 1852) F Cylindera (Eugrapha) contorta contorta (Fischer, 1828).
China (Shanxi, Heilongjiang, Gansu, Qinghai, Beijing, Hebei, Hubei, Sichuan), Russia, Mongolia.
Cicindela japanensis hamifasciata:
Cicindela hybrida transbaicalica:
Cicindela (Cicindela) transbaicalica hamifasciata:
Heilongjiang (
Beijing, Shunyi, leg. unknown, 13.v.1964, 2 males, 3 females (
China (Beijing, Heilongjiang, Jilin, Inner Mongolia, Liaoning, Gansu, Xinjiang, Shaanxi, Shanxi, Anhui, Henan, Hebei, Hubei, Jiangsu, Shandong, Sichuan, Qinghai, ?Fujian, ?Guangdong, ?Zhejiang, ?Yunnan), North Korea, South Korea, Russia.
New provincial record for Beijing.
Cicindela separata:
Cosmodela separata:
Shanxi (
Guangxi, Jinxiu, Tongmu Town, leg. J.C. Huang, 4.vii.1981, 2 females (
China (Guangxi, Guizhou, Shanxi, Jiangsu, Shanghai, Zhejiang, Henan, Anhui, Hubei, Hunan, Jiangxi, Fujian, Yunnan), Vietnam.
New provincial records for Guangxi and Guizhou.
Subgenus Cylindera Westwood, 1831
Cicindela obliquefasciata:
Cylindera (Cylindera) obliquefasciata obliquefasciata:
Heilongjiang (
Beijing, Haidian, Fragrant Hills Park, leg. unknown, 16.vii.1962, 2 males, 3 females (
China (Heilongjiang, Jilin, Inner Mongolia, Liaoning, Beijing, Hebei, Gansu, Qinghai, Xinjiang, Shanxi, Shandong, Henan), Russia.
Cicindela albopunctata:
Cylindera (Eriodera) albopunctata:
Sichuan (
Sichuan, Yanyuan, Jinhe, 1200 m, leg. D.J. Liu, 29.vi.1984, 1 female (
China (Sichuan, Yunnan, Xizang), Pakistan, Nepal, Bhutan, India, Vietnam.
Cylindera (Eugrapha) contorta contorta:
Gansu (
Inner Mongolia, Alxa League, Alxa Right Banner, leg. Y.C. Lv, 12.vii.1986, 1 male, 1 female (
China (Inner Mongolia, Gansu, Qinghai, Xinjiang), Ukraine, Romania, Moldova, Iran, Georgia, Azerbaijan, Kazakhstan, Uzbekistan, Turkmenistan, Tadzhikistan, Afghanistan, Russia, Mongolia.
Cicindela elisae:
Cylindera (Eugrapha) elisae elisae:
Heilongjiang (
Habitus photographs A Cylindera (Eugrapha) elisae elisae (Motschulsky, 1859) B Cylindera (Eugrapha) sublacerata vicaria (Semenov, 1895) C Cylindera (Ifasina) decolorata (Horn, 1907) D Cylindera (Ifasina) lesnei (Babault, 1923) E Cylindera (Ifasina) sikhimensis (Mandl, 1982) F Lophyra (Spilodia) lineifrons (Chaudoir, 1865). Scale bars: 5 mm.
Heilongjiang, Harbin, Hadeng (哈灯, hand-written label), leg. unknown, 13.vii.1963, 2 males, 1 female (NEFU).
Distribution maps. Green indicates records with previously known distribution based on published data, blue indicates new records, red stars indicate the collection sites of the examined specimens A Cylindera (Eugrapha) elisae elisae (Motschulsky, 1859) B Cylindera (Eugrapha) sublacerata vicaria (Semenov, 1895) C Cylindera (Ifasina) decolorata (Horn, 1907) D Cylindera (Ifasina) kaleea kaleea (Bates, 1866) E Cylindera (Ifasina) lesnei (Babault, 1923) F Cylindera (Ifasina) sikhimensis (Mandl, 1982).
China (Heilongjiang, Jilin, Inner Mongolia, Liaoning, Hebei, Shandong, Beijing, Shanxi, Gansu, Xinjiang, Anhui, Henan, Hubei, Hunan, Zhejiang, Guangdong, Hainan, Guangxi, Fujian, Shanghai, Jiangsu, Jiangxi, Qinghai, Sichuan, Yunnan, Xizang, Hong Kong, Taiwan), North Korea, South Korea, Vietnam, Russia, Mongolia.
Cicindela sublacerata:
Cicindela (Eugrapha) sublacerata:
Cylindera (Eugrapha) sublacerata vicaria:
Xinjiang (
Xinjiang, Changji, Fukang Desert Ecological System Observatary, 44°17'31"N, 87°56'3"E, 474 m, light trap, leg. Y. Liu, 11.vi.2007, 1 female (
China (Xinjiang), Mongolia.
We provide additional specimen data for this species from Xinjiang.
Cicindela psilica decolorata:
Cylindera (Ifasina) decolorate:
Fujian (
Guangxi, Huaping Nature Reserve, Mount Tianping, leg. J.K. Yang, 5.vi.1963, 1 male, 1 female (
China (Guangxi, Fujian, Guangdong, Sichuan, Guizhou, Yunnan), Vietnam.
New provincial record for Guangxi.
Cicindela kaleea:
Cylindera (Ifasina) kaleea kaleea:
Beijing (
Anhui, Tiantangzhai, Tudiling Bridge, 31°30'23"N, 116°09'07"E, 635 m, leg. unknown, 24.ix.2021, 1 male (
China (Anhui, Beijing, Hebei, Shaanxi, Gansu, Shandong, Jiangsu, Shanghai, Zhejiang, Jiangxi, Fujian, Henan, Hubei, Hunan, Guangdong, Guangxi, Sichuan, Guizhou, Yunnan, Xizang, Hong Kong, Taiwan), India, Myanmar, Thailand, Laos, Vietnam.
New provincial record for Anhui.
Cicindela lesnei: Babaule 1923: 7.
Cylindera (Ifasina) lesnei:
Guangdong (
Hunan, Huitong, Raochong Village, 26°51'23"N, 109°50'43"E, 650 m, leg. H.B. Liang, 21.vi.2015, 1 female (
China (Hunan, Guangdong), Vietnam.
New provincial record for Hunan.
Cicindela (Ifasina) discrete sikhimensis:
Cylindera (Ifasina) sikhimensis:
Xizang, Motuo, 2 km SE of Beibeng, 29°14'8"N, 95°9'31"E, 843 m, light trap, leg. H.B. Liang, 29.vii.2012, 4 males, 9 females (
China (Xizang), India, Myanmar.
New state record for China and new provincial record for Xizang.
Subgenus Spilodia Rivalier, 1961
Cicindela lineifrons:
Lophyra (Spilodia) lineifrons:
Yunnan (
Habitus photographs A Lophyra (Spilodia) striolata dorsolineolata (Chevrolat, 1845) B Heptodonta pulchella (Hope, 1831) C Pronyssiformia excoffieri (Fairmaire, 1897) D Therates biserratus Tan, Mo & Liang, 1991 E Therates guangdongensis Wiesner, 2016 F Therates hunanensis Matalin & Wiesner, 2023. Scale bars: 5 mm.
Guangxi, Ningming, Longrui, 180 m, leg. F.S. Li, 18.v.1984, 1 female (
China (Guangxi, Yunnan), Nepal, Bangladesh, India, Myanmar, Thailand, Cambodia, Laos, Vietnam, Malaysia.
New provincial record for Guangxi.
Cicindela dorsolineolata:
Lophyra (Spilodia) striolata dorsolineolata:
Beijing (
Distribution maps. Green indicates records with previously known distribution based on published data, blue indicates new records, red stars indicate the collection sites of the examined specimens A Lophyra (Spilodia) lineifrons (Chaudoir, 1865) B Lophyra (Spilodia) striolata dorsolineolata (Chevrolat, 1845) C Heptodonta pulchella (Hope, 1831) D Pronyssiformia excoffieri (Fairmaire, 1897) E Therates biserratus Tan, Mo & Liang, 1991 F Therates guangdongensis Wiesner, 2016.
Xizang, Beibeng, 2 km from Highway Gelin, 29°14'56"N, 95°11'20"E, 1013 m, leg. J.W. Jiang, 29.vii.2019, 3 males, 5 females (
China (Xizang, Beijing, Hebei, Shandong, Zhejiang, Jiangsu, Anhui, Jiangxi, Fujian, Henan, Hubei, Hunan, Guangdong, Guangxi, Hainan, Yunnan, Guizhou, Sichuan, Taiwan), Japan, Vietnam, Indonesia, Philippines.
New provincial record for Xizang.
Genus Heptodonta Hope, 1838
Cicindela pulchella:
Heptodonta pulchella:
Fujian (
Hainan, Jianfengling, Roadside of Tianchi, 18°43'44"N, 108°53'9"E, 1000 m, leg. H.B. Liang, 4.v.2007, 1 male, 2 females (
China (Hainan, Fujian, Guangdong, Guangxi, Yunnan, Sichuan, Xizang, Macao), Nepal, India, Myanmar, Thailand, Laos, Vietnam.
New provincial record for Hainan.
Cicindela excoffieri:
Pronyssiformia excoffieri:
Fujian (
Guizhou, Mount Fanjing, Huguosi, 27°54'44"N, 108°38'37"E, 1350 m, leg. Q.Z. Song, 3.viii.2001, 1 female (
China (Guizhou, Fujian, Hubei, Sichuan, Yunnan).
New provincial record for Guizhou.
Genus Therates Latreille, 1816
Therates biserratus:
Guizhou (Jiangkou; Yinjiang) (
Guangxi, Jinxiu, Luoxiang, 400 m, leg. D.C. Yuan, 14.v.1999, 1 male (
China (Guangxi, Jiangxi, Guizhou).
New provincial record for Guangxi and Jiangxi.
Therates guangdongensis:
Guangdong (Yinnah Shan) (
Hainan, Baisha County, Nanmaola, leg. X.L. Huang, 13.v.2009, 1 male (
China (Guangdong, Hainan).
New provincial record for Hainan.
Therates hunanensis:
Hunan (Dong’an; Chengbu) (
Chongqing, Mount Simian, 28°35–37'N, 106°23–24'E, 1190 m, leg. Z.H. Yang, 5.vii.2008, 1 male (
China (Chongqing, Hunan).
New provincial record for Chongqing.
Therates probsti:
Xizang, Motuo, Yarang Hydropower Station, 29°15'54"N, 95°14'43"E, 850 m, leg. H.B. Liang, 1.viii.2019, 1 female (
Habitus photographs A Therates probsti Wiesner, 1988 B Therates turnai Wiesner, 2015 C Neocollyris (Isocollyris) grandivadosa (Horn, 1935) D Neocollyris (Neocollyris) saphyrina (Chaudoir, 1850) E Neocollyris (Pachycollyris) bicolor (Horn, 1902) F Neocollyris (Pachycollyris) mouhotii nagaii Naviaux & Sawada, 1992. Scale bars: 5 mm.
China (Xizang), Laos, Vietnam.
Distribution maps. Green indicates records with previously known distribution based on published data, blue indicates new records, red stars indicate the collection sites of the examined specimens A Therates hunanensis Matalin & Wiesner, 2023 B Therates probsti Wiesner, 1988 C Therates turnai Wiesner, 2015 D Neocollyris (Isocollyris) grandivadosa (Horn, 1935) E Neocollyris (Neocollyris) saphyrina (Chaudoir, 1850) F Neocollyris (Pachycollyris) bicolor (Horn, 1902).
New state record for China and new provincial record for Xizang.
Therates turnai:
Guizhou (Xianheping) (
Sichuan, Mount Emei, Xixinsuo, 29°34'30"N, 103°22'26"E, 1300 m, day collecting, leg. H.B. Liang, 10–15.viii.2012, 1 male (
China (Guizhou, Sichuan).
New provincial record for Sichuan.
Subtribe Collyridini Brullé, 1834
Genus Neocollyris Horn, 1901
Subgenus Isocollyris Naviaux, 1994
Collyris aureofusca grandi-vadosa:
Neocollyris (Isocollyris) grandivadosa:
Guangxi, Napo County, 23°43'33"N, 106°49'30"E, 111 m, hand collected, leg. Y. Wang, 16.v.2021, 1 male, 1 female (
China (Guangxi), Vietnam.
New state record for China and new provincial record for Guangxi.
Neocollyris (Neocollyris) saphyrina:
?Yunnan (
Xizang, Shigatse, Gyirong, 2400 m, leg. F.S. Huang, 22.vii.1975, 1 female (
China (Xizang, ?Yunnan, ?Sichuan), India, Myanmar, Laos, Nepal, Bhutan, Bangladesh, Thailand, Indonesia.
Gyirong borders Nepal, from where N. saphyrina was previously known.
Collyris bicolor:
Neocollyris (Pachycollyris) bicolor:
Guangxi (
Hainan, Wuzhishan, Fanyang Town, leg. G. Ros, 1.vii.1956, 1 female (
China (Hainan, Guangxi, Yunnan), Laos, Vietnam.
New provincial record for Hainan.
Neocollyris mouhotii nagaii:
Neocollyris (Pachycollyris) mouhotii nagaii:
Hainan, Baisha County, Yinggeling, 18°59'28"N, 109°20'18"E, leg. M.Y. Lin, 14.vi.2010, 1 female (
China (Hainan), Vietnam.
New state record for China and new provincial record for Hainan.
Neocollyris sawadai:
Yunnan, Pu’er, Simao, day collecting, leg. H.L. Han and M.J. Qi, 15–19.vii.2009, 1 female (NEFU).
China (Yunnan), Vietnam.
Distribution maps. Green indicates records with previously known distribution based on published data, blue indicates new records, red stars indicate the collection sites of the examined specimens A Neocollyris (Pachycollyris) mouhotii nagaii Naviaux & Sawada, 1992 B Neocollyris (Pachycollyris) sawadai Naviaux, 1991 C Neocollyris (Pachycollyris) strangulata Naviaux, 1991 D Neocollyris (Pachycollyris) tricolor Naviaux, 1991 E Neocollyris (Pachycollyris) vitalisi (Horn, 1924).
New state record for China and new provincial record for Yunnan.
Neocollyris strangulata:
Neocollyris (Pachycollyris) strangulata:
Guangxi, Jingxi City, Diding Nature Reserve, 23°6'47"N, 105°58'40"E, leg. S.Y. Zhou and J.H. Huang, 9.viii.2010, 1 female (
China (Guangxi), Laos, Vietnam.
New state record for China and new provincial record for Guangxi.
Neocollyris tricolor:
Neocollyris (Pachycollyris) tricolor:
Yunnan (Lincang) (
Xizang, Beibeng, 2 km from Highway Gelin, 29°14'56"N, 95°11'20"E, 1013 m, leg. J.W. Jiang, 29.vii.2019, 1 female (
China (Xizang, Yunnan), Myanmar, Thailand, Laos, Vietnam.
New provincial record for Xizang.
Neocollyris (Pachycollyris) feai vitalisi:
Neocollyris (Pachycollyris) vitalisi:
Xizang (
Yunnan, Mount Huanglian, Beiluo, 22°44'8"N, 102°18'27"E, 1277 m, day collecting, leg. G.Z. Zhong and L.K. Zhang, 09–15.IX.2016, 1 female (
China (Yunnan, Xizang), Myanmar, Thailand, Laos, Vietnam.
New provincial record for Yunnan.
During our investigation of Chinese collections, we encountered a single specimen of Cicindela (Cicindela) japana Motschulsky, 1858 [Anhui, China = CHINE, Prov. ANHWEI-printed label] (Fig.
With the new records presented here, the tiger beetle fauna of China now includes 208 species, 71 (34%) of which are endemic. Five of the new records reported here are from the Chayu and Motuo, regions in the southeast of Xizang. This area is located at the junction of the Hengduan Mountains and the Himalayas and is surrounded by many biogeographical barriers. These five species were previously known from adjacent regions and countries, including Yunan Province, Sichuan Province, Myanmar, and India, from which they may have dispersed. More research on the tiger beetle diversity in this region, the southeast of Xizang, is needed and has great potential to broaden our understanding of the biogeography of tiger beetles within China.
Recently,
Furthermore, we report new provincial records of Therates biserratus Tan, Mo & Liang, 1991 and Therates hunanensis Matalin & Wiesner, 2023 in China. Therates biserratus has been included in the List of Key Protected Wild Animals in China since 2003, although it has not been considered as a separate species for a long time. According to the known distribution, T. biserratus may indeed be widely distributed in Nan Ling Area (24°00'–26°30'N, 110°–116°E). This may ultimately lead to reconsideration of the taxonomic status between these two species. Due to the limited number of specimens included in this and other studies (
To date, taxonomic studies on tiger beetles from Guangxi and Hainan are relatively scarce (
Many of the specimens reported here are old specimens that have not been previously identified. There are likely numerous unidentified tiger beetle specimens in other institutions and museums in China, which hold considerable yet untapped data. We encourage others to recognize and utilize this potential and are willing to help in identification.
We are grateful to Huilin Han (NEFU), Hongbin Liang (
The authors have declared that no competing interests exist.
No ethical statement was reported.
No funding was reported.
Conceptualization: KYW. Formal analysis: JW, FAB. Funding acquisition: CDL. Investigation: KYW. Resources: CDL. Supervision: CDL. Visualization: KYW. Writing - original draft: KYW. Writing - review and editing: FAB, JW.
Ke-Yi Wang https://orcid.org/0009-0009-8915-9563
Fabian A. Boetzl https://orcid.org/0000-0001-5121-3370
Jürgen Wiesner https://orcid.org/0000-0002-7824-166X
All of the data that support the findings of this study are available in the main text.