Yang Luo1, Thierry Bourgoin2, Jia-Lin Zhang1, Ji-Nian Feng1. 1. Key Laboratory of Plant Protection Resources and Pest Management, Ministry of Education, Entomological Museum, College of Plant Protection, Northwest A&F University, Yangling, Shaanxi 712100, China, Yangling, China Key Laboratory of Plant Protection Resources and Pest Management, Ministry of Education, Entomological Museum, College of Plant Protection, Northwest A&F University, Yangling, Shaanxi 712100, China Yangling China. 2. Institut de Systématique, Évolution, Biodiversité, ISYEB-UMR 7205, MNHN-CNRS-Sorbonne Université-EPHE-Univ. Antilles, Muséum national d'Histoire naturelle, CP 50, 57 rue Cuvier, F-75005, Paris, France Institut de Systématique, Évolution, Biodiversité, ISYEB-UMR 7205, MNHN-CNRS-Sorbonne Université-EPHE-Univ. Antilles, Muséum national d'Histoire naturelle, CP 50, 57 rue Cuvier, F-75005 Paris France.
Abstract
BACKGROUND: Cixiidae are small strictly phytophagous hemipteran insects worldwide distributed. Ecology and systematics of Chinese fauna remains poorly investigated. For instance, does their distribution follows the patterns of biogeogaphical distribution established for their host plants or other related-taxa because they are all obligatory phytophagous taxa? Do they follow the usual distributional Chinese realms and boundaries already recognized? Which zoogeographical Chinese regions and connections between them do they depict. To investigate these issues, we provide here a referenced and comprehensive checklist of the 250 cixiid species currently reported from China (77 new records), with their precise distribution at the regional level. In the 8 Chinese main zoogeographical regions usually recognized and 2 adjacent areas, we analyzed further their diversity at the tribal, generic, and specific levels using a non-metric multidimensional scaling and an unweighted pairwise group analysis using an arithmetic mean cluster analyses. The observed distribution patterns shown that an intercalary Sino-Japanese realm is recognisable between the Palaearctic and Oriental realms. At the regional level, the South China region clusters more closely with the Southwest, Central and North China regions. Taiwan, clearly separated from the South China region and mainland China, is more closely related to the Qinghai-Tibet region and Indochina countries. Although Central and South China regions remain close to each other, the Qinghai-Tibet region appears singularly different. NEW INFORMATION: An updated checklist of the 250 Cixiidae species, known to occur in China and counting for 10% of the Chinese planthopper fauna, is presented based on literature, recent collections, and museum records. More than 400 records distributed among the 28 provinces and 8 regions in China are extensively provided, including 77 new records. Of these, more than 80% of the species (205 species, 82%) have been only reported from China, and most of them are endemic species, which could reflects the great diversity degree of the Chinese regions and local biotypes highlights the uniqueness of this fauna. These species are found in 8 Chinese zoogeographical regions: The Taiwan region is the most diversified with 161 species and the highest rate of endemic species (69.57%), followed by South China (78 species, 17.95%), Central China (60 species, 33.33%), Southwest China (43 species, 39.53%), North China (29 species, 34.48%), Qinghai-Tibet region (10 species, 20%), Northeast China (8 species, 12.5%), and 5 species found in the Inner Mongolia-Xinjiang region that are not endemic ones. Endemism was analyzed for each region and repeated for species distribution patterns across them, 9 being bi-regionally and tri-regionally distributed. The South China-Taiwan pattern is the most richest one, followed by the Central-South China-Taiwan pattern. Semonini and Pentastirini tribes are widespread among all the zoological regions, representing respectively 21.20% and 17.20% of all the species, while Cixiini being is the most common tribe with 45.20%, remains absent from the North-Eastern China region. Andini with only 5.20% of the species is distributed in the Sino-Japanese - Oriental Region; Eucarpini (6.40%) and Borysthenini (2.00%) are mainly concentrated in the south of the Qingling Mountain-Huai River. The remaining four tribes, Bennini (0.40%), Briixini (0.80%), Oecleini (1.20%) and Stenophlepsiini (0.40%) are relatively rare and restricted to Taiwan. At the generic level, Kuvera (7.2%) is the most widely distributed genus in China while Cixius, Betacixius, Kuvera, Oecleopsis and Andes are the more diversified. One genus (Oliparisca) is distributed only in the Tibet region, while 10 genera are distributed only in the Taiwan region. In addition, nearly half of the genera (16 genera, 48.48%) are distributed south of the Palearctic/Oriental boundary. A non-metric multidimensional scaling and an unweighted pairwise group method analysis using arithmetic mean clustering based on the Jaccard similarity coefficient matrix support a Palaearctic/Sino-Japanese boundary and a South China region closer to the Southwest, Central and North China regions. The Taiwan region appears clearly separated from the South China region and to mainland China, and more closely related to the Qinghai-Tibet region and Indochina countries. The Central and South China regions appear close to each other, but the Qinghai-Tibet region is singularly isolated. Yang Luo, Thierry Bourgoin, Jia-Lin Zhang, Ji-Nian Feng.
BACKGROUND: Cixiidae are small strictly phytophagous hemipteran insects worldwide distributed. Ecology and systematics of Chinese fauna remains poorly investigated. For instance, does their distribution follows the patterns of biogeogaphical distribution established for their host plants or other related-taxa because they are all obligatory phytophagous taxa? Do they follow the usual distributional Chinese realms and boundaries already recognized? Which zoogeographical Chinese regions and connections between them do they depict. To investigate these issues, we provide here a referenced and comprehensive checklist of the 250 cixiid species currently reported from China (77 new records), with their precise distribution at the regional level. In the 8 Chinese main zoogeographical regions usually recognized and 2 adjacent areas, we analyzed further their diversity at the tribal, generic, and specific levels using a non-metric multidimensional scaling and an unweighted pairwise group analysis using an arithmetic mean cluster analyses. The observed distribution patterns shown that an intercalary Sino-Japanese realm is recognisable between the Palaearctic and Oriental realms. At the regional level, the South China region clusters more closely with the Southwest, Central and North China regions. Taiwan, clearly separated from the South China region and mainland China, is more closely related to the Qinghai-Tibet region and Indochina countries. Although Central and South China regions remain close to each other, the Qinghai-Tibet region appears singularly different. NEW INFORMATION: An updated checklist of the 250 Cixiidae species, known to occur in China and counting for 10% of the Chinese planthopper fauna, is presented based on literature, recent collections, and museum records. More than 400 records distributed among the 28 provinces and 8 regions in China are extensively provided, including 77 new records. Of these, more than 80% of the species (205 species, 82%) have been only reported from China, and most of them are endemic species, which could reflects the great diversity degree of the Chinese regions and local biotypes highlights the uniqueness of this fauna. These species are found in 8 Chinese zoogeographical regions: The Taiwan region is the most diversified with 161 species and the highest rate of endemic species (69.57%), followed by South China (78 species, 17.95%), Central China (60 species, 33.33%), Southwest China (43 species, 39.53%), North China (29 species, 34.48%), Qinghai-Tibet region (10 species, 20%), Northeast China (8 species, 12.5%), and 5 species found in the Inner Mongolia-Xinjiang region that are not endemic ones. Endemism was analyzed for each region and repeated for species distribution patterns across them, 9 being bi-regionally and tri-regionally distributed. The South China-Taiwan pattern is the most richest one, followed by the Central-South China-Taiwan pattern. Semonini and Pentastirini tribes are widespread among all the zoological regions, representing respectively 21.20% and 17.20% of all the species, while Cixiini being is the most common tribe with 45.20%, remains absent from the North-Eastern China region. Andini with only 5.20% of the species is distributed in the Sino-Japanese - Oriental Region; Eucarpini (6.40%) and Borysthenini (2.00%) are mainly concentrated in the south of the Qingling Mountain-Huai River. The remaining four tribes, Bennini (0.40%), Briixini (0.80%), Oecleini (1.20%) and Stenophlepsiini (0.40%) are relatively rare and restricted to Taiwan. At the generic level, Kuvera (7.2%) is the most widely distributed genus in China while Cixius, Betacixius, Kuvera, Oecleopsis and Andes are the more diversified. One genus (Oliparisca) is distributed only in the Tibet region, while 10 genera are distributed only in the Taiwan region. In addition, nearly half of the genera (16 genera, 48.48%) are distributed south of the Palearctic/Oriental boundary. A non-metric multidimensional scaling and an unweighted pairwise group method analysis using arithmetic mean clustering based on the Jaccard similarity coefficient matrix support a Palaearctic/Sino-Japanese boundary and a South China region closer to the Southwest, Central and North China regions. The Taiwan region appears clearly separated from the South China region and to mainland China, and more closely related to the Qinghai-Tibet region and Indochina countries. The Central and South China regions appear close to each other, but the Qinghai-Tibet region is singularly isolated. Yang Luo, Thierry Bourgoin, Jia-Lin Zhang, Ji-Nian Feng.
Entities:
Keywords:
Checklist; China; Cixiidae; distribution; endemism; species richness; zoogeography region
China covers an area of 9,634,057 km2, encompassing a area of entire Europe, and spans nearly 50 degrees of latitude from north to south, and more than 60 degrees of longitude from east to west in a world-renowned monsoon region (National Bureau of Statistics of the People’s Republic of China, http://data.stats.gov.cn). Most regions have cold, dry winters and warm, rainy summers, but in combination with the varying topography and terrain conditions, the climate is actually very complex and locally diverse with a wide variety of temperature zones and precipitation gradients (Ren and Wen 2011). Most regions are located in the temperate zone (semi-tropical, warm, mid-range, and cold-temperatures). A small portion of the country is in the tropics and plateau climate zone (the Qinghai-Tibet plateau temperate zone), and northern regions are close to the boreal zone (Jiang 2017). Annual precipitation decreases from the rain-forest of the southeast coast to the Gobi Desert in the northwestern interior (Jiang 2017). An arid humidity zone covers about 31% of the land area (mainly in northwest China). A semi-arid zone covers 22%, a semi-humid zone covers 15%, and the humid zone (32%) is located primarily in the southeast of China (Ge et al. 2013). Geological complexity of China is also significant, particularly with the uplift of the Qinhai-Tibet Plateau, which occurred in the middle of the Eocene era (45-38 Ma) (Zhou et al. 2018). When this complexity is combined with the monsoonal climate evolution, it has created strongly diversified biotopes, isolated by biogeographical barriers that manage dispersal pathways for species, providing new ecological niches, which has driven the recent evolution of plants and animal diversity (Favre et al. 2015, Liu et al. 2017).From a biogeographical point of view, China is usually divided in two parts, the Palearctic realm in the north, and the Oriental one in the south (Sclater 1858, Wallace 1876, Morrone 2015, He et al. 2017). From a zoological perspective, Holt et al. (2013) recently recognized an additional Sino-Japanese realm ranging from west of Tibet to the east of the Japanese archipelago standing between them. Accordingly, three main biogeographical lines cross China (Fig.1): the Palaearctic/Sino-Japanese boundary at about 40–41N, the Palearctic/Oriental line that follows the Qingling Mountain-Huai River, around 32–34N, and the Sino-Japanese/Oriental boundary at 24–25N in Southeastern China.The family Spinola, 1839 (: ), is a numerous and diverse taxon with a world-wide distribution (Holzinger et al. 2002, Bourgoin 2021). It comprises 18.6% of the currently known planthopper species (Bourgoin 2021), and is the largest family of the group. Classical taxonomy has divided the into 3 subfamilies: Emeljanov, 1989, Muir, 1923 and Spinola, 1839 (Holzinger et al. 2002). However, recent phylogenetical analyses have shown that these divisions remain artificial and three main lineages should better reflect of evolution of the family: an oecleinian lineage (including ), a cixiinian lineage and a pentastirinian lineage (including ) (Luo et al. 2021). Therefore, without including the fossil taxa, are currently divided into 18 tribes, 250 genera, and 2600 species (Bourgoin 2021).nymphs usually live underground and feed on plant rootlets, whereas the adults feed on the above ground phloem tissues of woody or plants and ferns (Wilson 1994, Wheeler 2003), predominaly on (9.2%), (7.8%), (6.7%), (6.5%), (5.1%) and (5.1%) in Eudicots and on (8.3%) in Monocots (Bourgoin 2021). Several species are considered to be vectors of plant pathogen including viruses, phytoplasmas and other prokaryotic-like organisms (Wilson 2005).Although the are one of the larger planthopper families, little is known about their ecology, distribution and host plants. In China, knowledge of this fauna is still fragmented and an overall comprehensive study is lacking. The first contribution was by Melichar (1902) who described 2 genera with 5 species from western China. Matsumura (1914) published 'Die Cixiinen Japans', describing 14 genera and 30 species, mostly from Taiwan. Kato (1932) focused on Northeastern China taxa, and published one new species. The first checklist of from the China mainland was provided by Hu (1935), who listed 11 species in 5 genera, which was updated by Metcalf (1936) in his 'Catalogue of the ’. Since then, many new species have been added. Jacobi (1944) reported 5 new species from the Fujian province. Fennah (1956) added 6 genera and 17 species from South China. Hori (1982) described 3 new species from Taiwan. Chou et al. (1985) described 7 species in 4 genera in his "Economic Insect Fauna of China ()". Tsaur provided a series of important contributions to the fauna from Taiwan, describing 155 species in 20 genera (Tsaur and Lee 1987, Tsaur et al. 1988, Tsaur 1989a, Tsaur 1989b, Tsaur 1990a, Tsaur 1990b, Tsaur et al. 1991a, Tsaur et al. 1991b, Tsaur and Hsu 2003, Tsaur 2009). Since then, several papers describing new recent taxonomic discoveries have been published (Wang 1991, Wang 1992, Huang 1995, Hua 2000, Liang 2001, Liang 2005a, Liang 2005b, Guo and Wang 2007, Guo et al. 2009, Guo and Feng 2010, Zhang and Chen 2011a, Zhang and Chen 2011b, Zhang and Chen 2013a, Zhang and Chen 2013b, Ren et al. 2014, Xing and Chen 2014, Bai et al. 2015, Li et al. 2016, Zhi et al. 2017, Zhi et al. 2018a, Zhi et al. 2018b, Luo et al. 2019a, Luo et al. 2019b, Zhi et al. 2019, Zhi et al. 2020a, Zhi et al. 2020b, Zhi et al. 2021).All of these studies primarily focused on taxonomical treats, with limited ecological and geographical interpretations or evaluations. However, are obligatory phytophagous taxa and therefore directly linked to the distribution of their host plants (Attié et al. 2008). They are generally considered feeding on a variety of plants (Larivière 1999) but more precisley documented, they appears mostly oligiphagous or monphagous (Wilson et al. 1994, Bourgoin 2021). The planthopper and its host-plants are both patterned by the historical biogeography of the areas where they are distributed. How do follow the patterns of biogeogaphical distribution (major biological realms, biogeographical regions) already well established in China? Which boundaries can be identified for and at which taxonomical levels? The aim of this paper is to identify these correlations and to investigate how these zoogeographical regions are connected in China.This current paper provides the first distribution pattern of the Chinese following current Chinese zoogeographical regions recognized and updated species list of Chinese . Accordingly, the objectives of this paper are: (1) to compare species richness at the level of the Chinese zoogeographical regions and to document their distribution patterns and their endemism in each region, both at the tribal and generic level; (2) to investigate what biogeographical patterns the reflect: are they recognized effectively in a particular Sino-Japanese realm or a simple area of transition between the Palearctic and Oriental realms? (3) to provide a comprehensive species list of the from China.
Materials and methods
Eight Chinese zoogeographic regions, based on geographic, climatic, and vegetation characteristics (Gao et al. 2017, He et al. 2017), were used for the bio-geographical analyses: Northeast China, North China, Nei Mongol-Xinjiang, Qinghai-Tibet, South China, Central China, Southwest China and the Taiwan region (Fig. 1). Two other regions were added for countries adjacent to China: 1) a south China 'VM region' including Vietnam, Laos, Thailand, Cambodia, Myanmar, Bhutan, Bangladesh, as well as a small portion of India, and 2) a north East China 'Far East region' including a portion of Russia. The map (Fig. 1) was created using the National Earth System Science Data Sharing Infrastructure (http://www.geodata.cn).
Figure 1.
Map of zoogeographical regions of China and adjacent areas. Abbreviations: NEC, Northeast China; NC, North China; NX, Nei Mongol-Xinjiang; QT, Qinghai-Tibet; SWC, Southwest China; CC, Central China; SC, South China; TW, Taiwan; RFE, Russian Far East; VM, Vietnam, Laos, Thailand, Cambodia, Myanmar, Bhutan, Bangladesh and part of Indian.
The distribution matrix includes 253 Chinese species (of which 87 species were recorded from museums and the remaining species were recorded from the literature). Among them, 3 species: Kramer, 1981, Fieber, 1876, and (Olivier, 1791), were excluded from the analyses and checklist because we could not confirm their occurrence in China (no specimens information was found in our inspection of museum specimens in the collections) or because of uncertainties about where they were collected. 48 additional species (Suppl. material 1) from adjacent areas based on literature and FLOW (Bourgoin 2021) were added for the cluster analysis. The observed material information of checklist, as a formatted Excel spreadsheet, are provided here in the supplementary materials: Suppl. material 2. Figure 2 and 3 were generated using ArcGIS Version 10.8 statistical software (URL: https://desktop.arcgis.com/en/system-requirements/latest/arcgis-desktop-system-requirements.htm). The distribution information of the in China was imported into ArcGIS Version 10.8 software, the latitude and longitude of the distribution sites were set as the coordinate attribute elements, and the symbols in the map were set to different colors for distinguishing different genera of the tribes, and finally the maps of the distribution of the tribes and species were exported.Presence/absence matrices for species and for genera were built for each of the 10 OGUs (physiographical regions as operative geographical units, Crovello 1981). Similarity coefficients use binary data to measure association between OGU. On the basis of a review of similarity coefficients (Shi 1993), the Jaccard's coefficient in NTSYS Version 2.1 software (Rohlf 2000) was used according toLegendre and Legendre (1983) and Rohlf (2000). Clustering of OGUs using the UPGMA algorithm, UPGMA (an unweighted pairwise group method using arithmetic mean) was used to cluster similarities (Legendre and Legendre 1983). Based on the similarity of clustering results, Jaccard's coefficients were analyzed through nonmetric multidimensional scaling (NMDS) according to Kenkel and Orloci (1986).
Data resources
This publication follows the classical systematic classification based on Holzinger et al. (2002) and Emeljanov (2002) as synthetized and updated in Bourgoin (2021) and Luo et al. (2021). Fossil species are indicated by the symbol (†). The checklist contains information updated up to April, 2021 compiled from scientific papers, book chapters, conference abstracts, theses, and from the FLOW website (Bourgoin 2021). It also includes our own unpublished taxonomic data and original museum specimens information from the following institutions: Shanghai Entomological Museum C.A.S (SEM), Museum of China Agricultural University (CAU), Entomological Museum of Northwest A&F University (NWAFU), Museum of Chinese Academy of Forestry (CAF), Institute of Entomology, Guizhou University (GZU), Chongqing Normal University (CQNU) and Muséum National d'Histoire Naturelle (MNHN). Distribution sets were collected from the original sources with their original latitude and longitude information; Those lacking such information were approximed with the latitude and longitude coordinates of the corresponding administrative center.
New record: China: Hainan (Diaoluo Mountain).Spinola, 18399E5B213C-EC21-53E2-B747-2B9266786882Emeljanov, 2002C7C24FC8-F355-5C87-8845-4F4C57AAECC5Stål, 186689F4DF6B-2313-567F-B6C9-A37B060AB93A(Matsumura, 1914)9AC85C00-E651-5459-AA38-525D6DADFA62Matsumura, 1914: 432.|China: Fujian, Sichuan, Taiwan (Matsumura 1914).New record: China: Fujian (Wuyi Mountain).Fennah, 19780024AB39-4781-5128-80C0-FEA1B5CD49F2Fennah, 1978: 209.China: Yunnan; Malaysia: Kuala Lumpur, Kedah (Fennah 1978); Vietnam: Ninh Bình (Fennah 1978).New record: China: Yunnan (Menglun).Fennah, 195666EEBBE7-98FD-5990-867C-7D49129C5B4EFennah, 1956: 447.China: Zhejiang (Fennah 1956).Tsaur & Hsu, 1991C58CAA76-0D41-5664-84DA-0DEBBD4FB6CCTsaur & Hsu in Tsaur et al., 1991a: 72.China: Zhejiang, Taiwan (Tsaur et al. 1991a).(Uhler, 1896)9B684D7D-4B4F-54B7-8B9A-F867FE65AD74Uhler, 1896: 280.|China: Beijing (Liang 2005b), Henan, Jiangsu, Zhejiang, Guangxi, Guizhuo; Japan: Hokkaido, Honshu, Shikoku, Kyushu, Tsushima Island (Palaearctic) (Hayashi and Fujinuma 2016); Russia: Far East.New record: China: Jiangsu (Suzhou), Zhejiang (Taishun).Fennah, 19560E353325-7094-52D9-B5B5-C539238C781EFennah, 1956: 446.| Zhang, 2008: 33.China: Beijing, Henan, Hubei (Fennah 1956), Guizhou.New record: China: Beijing (Mentougou), Henan (Huixian), Hubei (Lichuan).Tsaur & Hsu, 1991E33759D0-0393-507D-90B5-50ADC287AB45Tsaur & Hsu in Tsaur et al., 1991a: 70.China: Guangxi, Tibet, Taiwan (Tsaur et al. 1991a).New record: China: Guangxi (Jinxiu, Longsheng), Tibet (Motuo).Fennah, 1956720922F3-6F07-5EE5-9B81-3EBEB320F675Fennah, 1956: 445.China: Hong Kong (Fennah 1956).Fennah, 19787698DE80-07B3-51BF-9357-9DA2DA770498Fennah, 1978: 208.China: Guizhou, Zhejiang; Vietnam: Ninh Bình (Fennah 1978).New record: China: Zhejiang (Fengyang Mountain).Fennach, 195657BB74AA-F3FC-5EF0-A376-857C228F2DBEFennach, 1956: 444.| Zhang, 2008: 38.China: Guangdong, Sichuan (Fennah 1956).Emeljanov & Hayashi, 2007120AAE2E-AC65-59B3-A453-5B17DE075BA1Zhi & Chen, 2018B0494B80-72AB-567F-9236-B5B172198D7EZhi & Chen in Zhi et al., 2018b: 57.China: Yunnan (Zhi et al. 2018b).Zhi & Chen, 2018A4A12AE6-C39B-5BE3-8CB3-3AF6743D8F32Zhi & Chen, in Zhi et al., 2018b: 60.China: Yunnan (Zhi et al. 2018b).Tsaur & Hsu, 1991E68BD9D9-FAAE-5C34-ADCE-D72030976C66Tsaur & Hsu in Tsaur et al., 1991a: 65.|China: Taiwan (Tsaur et al. 1991a).Metcalf, 193884BFA855-3303-5C5F-9206-31E41732A775Matsumura, 1938ABCCB77F-E995-5B39-9313-3249C2E63B72(Matsumura, 1938)DA5A57E8-56C2-5192-A719-885E735D66B2Matsumura, 1938: 152.|China: Taiwan (Tsaur 2009).Emeljanov, 2002336F0E07-09EF-565E-BA6C-E0FEB7F87020Stål, 185950004930-32B2-510F-9B2F-83BDFBBA2311Matsumura, 1914793EB166-9F6A-569B-9D64-952431BC1F74Matsumura, 1914: 433.China: Taiwan (Matsumura 1914).Van Stalle, 19833BFA30B4-B44D-55FC-880E-155DA38871FEVan Stalle, 1983: 272.China: Taiwan.Spinola, 1839E6EA5E4F-334B-552A-94DF-DF61D4A51544Tsaur & Hsu, 19910A7B2063-4DEA-50A9-90B1-3B8725938217Tsaur & Hsu, 1991B3FFF07B-EEFA-582A-93AA-5A80B4FD2439Tsaur & Hsu in Tsaur et al., 1991a: 19.China: Taiwan (Tsaur et al. 1991a).Tsaur & Hsu, 1991A8401FC3-6D09-5220-97F0-F48084260A53Tsaur & Hsu in Tsaur et al., 1991a: 12.China: Taiwan (Tsaur et al. 1991a).Tsaur & Hsu, 1991CFAE1F43-0443-5A7F-839F-84A5191AA2A3Tsaur & Hsu in Tsaur et al., 1991a: 17.China: Taiwan (Tsaur et al. 1991a).Tsaur & Hsu, 1991ADC7EF88-3DC9-5078-B205-FE5BD236B4CBTsaur & Hsu in Tsaur et al., 1991a: 9.China: Taiwan (Tsaur et al. 1991a).Tsaur & Hsu, 199142C5C12A-F647-54B2-96B0-02A17D2A0B52Tsaur & Hsu in Tsaur et al., 1991a: 9.China: Taiwan (Tsaur et al. 1991a).Tsaur & Hsu, 199100CE480F-E585-5CA7-9B54-D5231F94ADA7Tsaur & Hsu in Tsaur et al., 1991a: 14.China: Taiwan (Tsaur et al. 1991a).Tsaur & Hsu, 19911833FAE1-0666-5EC1-920E-D49754D6C0BATsaur & Hsu in Tsaur et al., 1991a: 14.China: Taiwan (Tsaur et al. 1991a).Latreille, 180463F8E04D-E8EC-5994-A9FF-C96B872A4F1ETsaur & Hsu, 1991F82D2B1F-56DD-519C-B332-5B1DC87CA661Tsaur & Hsu in Tsaur et al., 1991b: 199.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19911DE9C415-83D2-524E-95E8-3E49815585BBTsaur & Hsu in Tsaur et al., 1991b: 204.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991A53BE908-C0CE-5E10-BEEB-A4C36C71A9B1Tsaur & Hsu in Tsaur et al., 1991b: 239.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19919C8C06C6-E0A6-538A-A11B-6131AEF6E750Tsaur & Hsu in Tsaur et al., 1991b: 242.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199154DAD350-AFBA-5AA4-99B6-A1EDA4853CAATsaur & Hsu in Tsaur et al., 1991b: 266.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991C027BBDF-3194-54D5-BE52-550EA2518DEBTsaur & Hsu in Tsaur et al., 1991b: 260.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991A21D1D76-2E7D-53F4-B962-10172C19830CMatsumura, 1914: 386.| Tsaur et al., 1991b: 185.China: Zhejiang, Hainan, Taiwan (Tsaur et al. 1991b).New record: China: Zhejiang (Fengyang Mountain).Matsumura, 1914A4F985B9-6F8A-5828-8D27-32E73453CC6CMatsumura, 1914: 395.| Esaki, 1932: 1773.| Tsaur et al., 1991b: 175.China: Taiwan (Tsaur et al. 1991b), Hainan; Japan.New record: China: Hainan (Limuling).Tsaur & Hsu, 199171CC12A3-459B-53ED-8B1A-610D0475115BTsaur & Hsu in Tsaur et al., 1991b: 279.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991B7739556-840C-5FA6-A290-758760E95830Tsaur & Hsu in Tsaur et al., 1991b: 222.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19919E28851C-50AC-56D8-8C56-23CF6C1CC6F5Tsaur & Hsu in Tsaur et al., 1991b: 233.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991F70EA667-CCDC-5CC3-A682-BDA49F7C69C6Tsaur & Hsu in Tsaur et al., 1991b: 208.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19919C1AB30C-B3A9-5913-863C-B10EC5ABE256Tsaur & Hsu in Tsaur et al., 1991b: 275.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19916D37E211-4B8E-536B-B94B-64E94AD3305CTsaur & Hsu in Tsaur et al., 1991b: 236.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19917BF767BE-3089-530A-B914-D6983DD76AF3Tsaur & Hsu in Tsaur et al., 1991b: 225.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199162F98740-FD80-5391-ADBB-1D5D59F9A387Tsaur & Hsu in Tsaur et al., 1991b: 252.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19916B64193C-00A6-56C6-8A08-CD544558AA94Tsaur & Hsu in Tsaur et al., 1991b: 202.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19912E89B9FF-45E6-5F5D-A4CE-0A311C7496B8Tsaur & Hsu in Tsaur et al., 1991b: 219.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991C565BCEA-ED14-5535-9312-8FD157E683DCTsaur & Hsu in Tsaur et al., 1991b: 200.China: Guangxi, Zhejiang, Taiwan (Tsaur et al. 1991b).New record: China: Guangxi (Nanning), Zhejiang (Fengyang Mountain).Tsaur & Hsu, 19914D57F482-9BA4-573E-B697-337992640B15Tsaur & Hsu in Tsaur et al., 1991b: 183.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199138EF48CC-54A4-58EA-9027-4A2422E83CEBTsaur & Hsu in Tsaur et al., 1991b: 256.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199189078885-E950-5920-8BAA-36A7999F3789Tsaur & Hsu in Tsaur et al., 1991b: 300.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19912169A1BA-70B3-5912-A6F9-0C24B3B3FCC5Tsaur & Hsu in Tsaur et al., 1991b: 294.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199189BE279B-7B77-528E-90F3-C95F89EBF639Tsaur & Hsu in Tsaur et al., 1991b: 254.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991795BAA48-4D68-5314-BDC6-E4BCBC212E0ETsaur & Hsu in Tsaur et al., 1991b: 180.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199141AF7679-1851-5749-87F5-7D6F4E29147CTsaur & Hsu in Tsaur et al., 1991b: 241.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19911BF55B8B-42CC-5214-86AD-7B16086BAC3CTsaur & Hsu in Tsaur et al., 1991b: 286.China: Taiwan (Tsaur et al. 1991b).Li, Liu, Ren, Li & Yao, 20162B9B60C4-66FE-53FB-9EFD-C5D490D54301† Li, Liu, Ren, Li & Yao in Li et al., 2016: 2.China: Qinghai (Li et al. 2016).Fossil speciesTsaur & Hsu, 19918B386C62-4D09-548A-B867-DC7DE93FEA81Tsaur & Hsu in Tsaur et al., 1991b: 244.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991832A46BC-9E39-5D42-B76D-E904FD2F1306Tsaur & Hsu in Tsaur et al., 1991b: 206.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199128AC5137-D465-5958-986B-DE437A348BE2Tsaur & Hsu in Tsaur et al., 1991b: 180.China: Taiwan (Tsaur et al. 1991b).Matsumura, 19144FDBA5A1-8679-5125-A540-29A8B9BDD162Matsumura, 1914: 405.China: Shaanxi, Taiwan (Matsumura 1914).New record: China: Shaanxi (Hanzhong).Tsaur & Hsu, 1991CE2F1785-CAB0-5626-BC48-12A9307A83E3Tsaur & Hsu in Tsaur et al., 1991b: 285.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991617AC248-F0A1-5211-BC61-7778D7000949Tsaur & Hsu in Tsaur et al., 1991b: 228.China: Taiwan (Tsaur et al. 1991b).Fennah, 1956937C701D-3887-5CE9-9B7A-A6492E4B5F25Fennah, 1956: 449.China: Guangdong (Fennah 1956).Tsaur & Hsu, 19911D9DFCBC-67AE-5F49-B6C1-9E7AAC20CBD1Tsaur & Hsu in Tsaur et al., 1991b: 225China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991693B3781-428F-5BBF-BEF5-A26EBBBCB8B2Tsaur & Hsu in Tsaur et al., 1991b: 188.China: Taiwan (Tsaur et al. 1991b).Matsumura, 1914906EA96A-5DB2-5A3E-BC3A-2D7C4BEA9FE7Matsumura, 1914: 399| Tsaur et al., 1991b: 195.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19911C8DCE35-111D-502C-96D3-B510EB59FC6DTsaur & Hsu in Tsaur et al., 1991b: 256.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991C19B0B61-9836-5802-B8E7-1CF2059FC936Tsaur & Hsu in Tsaur et al., 1991b: 279.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19916EC1BDE8-C72C-5581-AE3A-E244D73E3612Tsaur & Hsu in Tsaur et al., 1991b: 212.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991E8542DF3-D523-5DFB-BA1D-38DF7AF8DFF4Tsaur & Hsu in Tsaur et al., 1991b: 219.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991EEDB50D3-ACCA-574F-B8D3-86FC1DDBB003Tsaur & Hsu in Tsaur et al., 1991b: 272.China: Taiwan (Tsaur et al. 1991b).Matsumura, 19149B56CBD4-47B2-5779-AD02-9EB9DE5DBDD7Matsumura, 1914: 401| Tsaur et al., 1991b: 301.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991C431752E-E30C-5787-A439-0AE84723D8C7Tsaur & Hsu in Tsaur et al., 1991b: 269.China: Taiwan (Tsaur et al. 1991b).Matsumura, 191446245028-261A-5C17-A1A3-65C394A16CD9Matsumura, 1914: 398.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991A6C62A6B-DFD0-513F-A06A-FD30895AE82ATsaur & Hsu in Tsaur et al., 1991b: 272.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199194FDE82A-B965-5F43-9CB4-F4DD0F64906ATsaur & Hsu in Tsaur et al., 1991b: 248.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991ABC7D7D4-15B2-58E0-A3F4-5D975E2C284FTsaur & Hsu in Tsaur et al., 1991b: 282.China: Zhejiang, Taiwan (Tsaur et al. 1991b).New record: China: Zhejiang (Feng Mountain).Tsaur & Hsu, 1991E8C80E02-88BD-5B5D-911E-7B17961A376BTsaur & Hsu in Tsaur et al., 1991b: 216.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991D7079E9D-E8C4-53D8-97C3-9D8410272F6ETsaur & Hsu in Tsaur et al., 1991b: 264.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199146E7B3B3-2910-57D1-B3E6-73804DC7C430Tsaur & Hsu in Tsaur et al., 1991b: 188.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199165875310-07BB-526C-ACAE-FE590576611DTsaur & Hsu in Tsaur et al., 1991b: 208.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991735383BE-2B54-5328-8206-0BA34A9C2D01Tsaur & Hsu in Tsaur et al., 1991b: 205.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991C666BBEA-8413-532F-AFF4-6B7A326171EFTsaur & Hsu in Tsaur et al., 1991b: 176.China: Fujian, Taiwan (Tsaur et al. 1991b).First record: China: Fujian (Wuyi Mountain).(Linné, 1758)9AE644EE-4940-5448-A85D-4D0230EFD460Linné, 1758: 437.|China: Ningxia; Algeria (Nast 1972, Holzinger et al. 2003) Austria (Nast 1972, Holzinger et al. 2003) Belgium (Nast 1972, Holzinger et al. 2003); Canada: Alberta, British Columbia, Manitoba, New Brunswick, Newfoundland, Nova Scotia, Ontario, Quebec, Saskatchewan (Bartlett et al. 2014) Czechoslovakia (Nast 1972, Holzinger et al. 2003); Denmark (Nast 1972, Holzinger et al. 2003); Finland (Nast 1972, Holzinger et al. 2003); France (Nast 1972, Holzinger et al. 2003); Germany (Nast 1972, Holzinger et al. 2003); Great Britain (Nast 1972, Holzinger et al. 2003); Hungary (Nast 1972, Holzinger et al. 2003); Italy (Nast 1972, Holzinger et al. 2003); Japan: Hokkaido, Honshu; Macedonia; Morocco; Netherlands; Norway; Poland; Romania; Russia; Serbia; Spain; Sweden; Switzerland; Tunisia; USA: Alaska, Arizona, California, Colorado, Connecticut, Delaware, Georgia, Idaho, Illinois, Indiana, Iowa, Kansas, Maine, Maryland, Massachusetts, Michigan, Minnesota, New Hampshire, New Jersey, New Mexico, New York, North Carolina, Ohio, Oregon, Pennsylvania, Rhode Island, South Dakota, Tennessee, Utah, Vermont, Virginia, Washington, Wisconsin (Bartlett et al. 2014).New record: China: Ningxia (Liupan Mountain).Matsumura, 1914F0889B2F-2742-5663-A300-E9FDBB654A89Matsumura, 1914: 401.| Jacobi, 1944: 14.| Schumacher, 1915: 131.China: Fujian, Taiwan (Matsumura 1914).Tsaur & Hsu, 19912511261C-2074-50E7-817B-930B4D5E5BF3Tsaur & Hsu in Tsaur et al., 1991b: 288.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19917398E183-14EE-568A-AD6D-8ECAF23D8133Tsaur & Hsu in Tsaur et al., 1991b: 247.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19919DBA15A9-AE2C-512B-8446-C5296328530ETsaur & Hsu in Tsaur et al., 1991b: 251.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19915C546A07-EA28-520F-9D6F-DB41933C8241Tsaur & Hsu in Tsaur et al., 1991b: 289.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991A9F3EF34-8C55-57F7-837E-048ECAE90CE4Tsaur & Hsu in Tsaur et al., 1991b: 236.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991F8E37213-BD8E-5E3B-B29D-59DC1D92658ETsaur & Hsu in Tsaur et al., 1991b: 291.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199157EB871F-C6FE-5514-8D75-01A15D228886Tsaur & Hsu in Tsaur et al., 1991b: 231.China: Taiwan (Tsaur et al. 1991b).Fennah, 1956FBE194C4-3DA8-5C39-AD5E-DAC6831CD4D2Fennah, 1956: 449 (China: Guangdong.Matsumura, 1914FD706C01-92B5-5A32-B396-C775FF4FC01FMatsumura, 1914: 405.China: Taiwan (Matsumura 1914).Tsaur & Hsu, 19915CFC8CEB-CF69-504C-9E56-609AE12A510ATsaur & Hsu in Tsaur et al., 1991b: 297.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991F6EA01D5-3227-5A05-95B1-37C7AFACD5A7Tsaur & Hsu in Tsaur et al., 1991b: 244.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991C103796E-3F3E-534F-870A-173919101B05Tsaur & Hsu in Tsaur et al., 1991b: 278.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19910CC83693-5BC7-5A1D-82C0-64D01DD9F810Tsaur & Hsu in Tsaur et al., 1991b: 249.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991B6A7A21A-10FA-5439-A136-FBA8976DE0C3Tsaur & Hsu in Tsaur et al., 1991b: 190.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199107F68428-C60D-5D7C-81A5-019B21D3D1E2Tsaur & Hsu in Tsaur et al., 1991b: 260.China: Taiwan (Tsaur et al. 1991b).Fennah, 1956B2F27C94-90D0-5E9A-894A-5FAA1A03B8F0Fennah, 1956: 450.| Tsaur et al., 1991b: 297.China: Anhui (Fennah 1956), Hunan, Henan, Taiwan (Tsaur et al. 1991b).New record: China: Hunan (Mang Mountain), Hunan (Huping Mountain), Anhui (Guniujiang), Henan (Yuhuang).Tsaur & Hsu, 199170B2127E-7F19-5BA3-B763-BA105F36BA0DTsaur & Hsu in Tsaur et al., 1991b: 263.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19914C534B50-5317-589D-BBCA-EE3D92BA53D7Tsaur & Hsu in Tsaur et al., 1991b: 214.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991068325B8-EDA2-5799-975A-A78D3DE69D34Tsaur & Hsu in Tsaur et al., 1991b: 266.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19912F84A3BE-1C71-5194-8AAC-93E5FDB9BE6ETsaur & Hsu in Tsaur et al., 1991b: 282.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19917D971480-F68E-522E-BE46-BC7682B9B33DTsaur & Hsu in Tsaur et al., 1991b: 186.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991F783E185-CD78-53EA-ABE8-05727E62899BTsaur & Hsu in Tsaur et al., 1991b: 291.China: Taiwan (Tsaur et al. 1991b).Matsumura, 19143F44CC3C-0E14-5A3A-B66B-EA6ADC32E02CMatsumura, 1914: 401| Tsaur et al., 1991b: 301.China: Taiwan (Matsumura 1914).Tsaur & Hsu, 1991C8C3354F-16C3-5AF8-A0C8-F4D573F670ECTsaur & Hsu in Tsaur et al., 1991b: 275.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991D1C2D33F-CC41-5773-99B1-FBB5A94FED80Tsaur & Hsu in Tsaur et al., 1991b: 294.China: Taiwan (Tsaur et al. 1991b).Matsumura, 19148D1A7769-442F-542A-9C85-201B110E22D2Matsumura, 1914: 398| Tsaur et al., 1991b: 195.China: Zhejiang, Taiwan (Tsaur et al. 1991b).New record: China: Zhejiang (Longwang Mountain).Tsaur & Hsu, 1991E9E2B57C-FCBB-5246-8997-706E4B713A11Tsaur & Hsu in Tsaur et al., 1991b: 229.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199104D99A90-775B-546C-B4E9-2B35692C02B8Tsaur & Hsu in Tsaur et al., 1991b: 259.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 19913D74EBD9-A204-5366-9385-87016D03EF23Tsaur & Hsu in Tsaur et al., 1991b: 233.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991CC9E81AE-9B36-5E27-B2DD-F27E88EF571DTsaur & Hsu in Tsaur et al., 1991b: 222.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 1991F86779CF-03FD-5E31-834E-8EB8FF5B55B5Tsaur & Hsu in Tsaur et al., 1991b: 216.China: Taiwan (Tsaur et al. 1991b).Matsumura, 1914AE8D9C25-4371-5ED4-B770-0680DE58F73DMatsumura, 1914: 403.China: Taiwan (Matsumura 1914).Tsaur & Hsu, 199115330322-B680-518B-B487-48CB2CAA6E43Tsaur & Hsu in Tsaur et al., 1991b: 211.China: Guangxi, Ningxia, Taiwan (Tsaur et al. 1991b).New record: China: Ningxia (Liupan Mountain), Guangxi (Huaping nature reserve).Tsaur & Hsu, 199147BD70F6-E135-551E-AF21-A4C0ACFB600BTsaur & Hsu in Tsaur et al., 1991b: 178.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199148F55DD6-70F0-5F83-BFBA-C33DAF1692A3Tsaur & Hsu in Tsaur et al., 1991b: 269.China: Taiwan (Tsaur et al. 1991b).Tsaur & Hsu, 199133CBE7AA-FA8E-5976-802E-2CFBD86C2EE2Tsaur & Hsu in Tsaur et al., 1991b: 192.China: Taiwan (Tsaur et al. 1991b).Wagner, 1939EF5165FF-EC18-529B-94FE-969DA492BDF4(Germar, 1818)9F418B54-257A-5A00-9884-D6DEE1614160Germar, 1818: 199.|China: Guangxi, Guizhou, Zhejiang; France (Ribaut and Lacroix 1958); Germany (Holzinger et al. 2003); UK (Holzinger et al. 2003); Iran: Kandovān (Mozaffarian and Wilson 2011); Netherlands (De Haas and Den Bieman 2018); Poland (Gebicki et al. 2013).New record: China: Guangxi (Huaping), Zhejiang (Hangzhou).Wagner, 193990E3DCFE-59E7-563B-9F31-4679FF4777BEVilbaste, 19688A5193E5-1BC3-51A2-B887-65D3CA2783E5Vilbaste, 1968: 5.| Anufriev & Emeljanov, 1988: 452.|China: Gansu.New record: China: Gansu (Wenxian).Tsaur & Hsu, 19913A91091B-82C9-57B2-B78E-601D14BEAD36Tsaur & Hsu, 1991DC6F69CE-D1AB-59FE-B628-1806A268ED1DTsaur & Hsu in Tsaur et al., 1991a: 25.China: Taiwan(Tsaur et al. 1991a).Matsumura, 19148EBE9C65-309F-5F97-B3F5-04E1C963CCAEMatsumura, 19147965E741-4BE1-5D63-8869-9FE072989844Matsumura, 1914: 394.| Schumacher, 1915: 131.| Fennah, 1956: 459.| Tsaur et al., 1991a: 3.| Liang, 2005: 429.| Orosz, 2013: 107.| Zhang & Chen, 2013b: 279.| Hayashi & Fujinuma, 2016: 325.China: Fujian, Hainan, Jiangxi (Zhang and Chen 2013a), Taiwan (Tsaur et al. 1991a); Japan: Kyushu; Vietnam (Hayashi and Fujinuma 2016).New record: China: Fujian (Chongan), Hainan (Jianfeng), Jiangxi (Wuyishan).Tsaur & Hsu, 199162A20272-C6C5-5F98-9370-4D8287C0D5CBTsaur & Hsu in Tsaur et al. 1991a: 5.| Orosz, 2013: 108.| Zhang & Chen, 2013b: 281.China: Guizhou, Sichuan, Yunnan, Zhejiang (Zhang and Chen 2013a), Taiwan (Tsaur et al. 1991a); Vietnam.New record: China: Guizhou (Luodian).Zhang & Chen, 20136FF32F2F-2935-581A-89F6-9629F8731AADZhang & Chen, 2013a: 283.| Orosz & Redei, 2016: 376.China: Guizhou (Zhang and Chen 2013a), Jiangxi, Taiwan (Tsaur et al. 1991a); Nepal: Ganesh Himal.Zhang & Chen, 2013917E8EAC-3D2A-54DD-8EB8-73EFECA23149Zhang & Chen, 2013a: 285.China: Yunnan (Zhang and Chen 2013a) .Tsaur & Hsu, 199104CF99CF-C06D-5829-A7EE-174190280F3ETsaur & Hsu, 1991E46700BA-100D-56EE-AE64-4DAE93EF1327Tsaur & Hsu in Tsaur et al., 1991a: 23.China: Taiwan (Tsaur et al. 1991a).Kocak & Kemal, 20097875DF93-D0AE-5BDC-9661-20FB930ED5F1Zhi & Chen, 201960BE95E6-7AFB-5A3A-AC97-F85272C0DDC3Zhi & Chen, 2019: 57.China: Guizhou, Hubei (Zhi et al. 2019).(Zhang & Chen, 2011)D9DDA4B9-2F07-5C23-8FF5-EE3C8EC609E0Zhang & Chen, 2011a: 61.|China: Guizhou (Zhang and Chen 2011a).Zhi & Chen, 20197ED7A33A-AFC8-57BF-8BCB-5E77C5E4EABCZhi & Chen, 2019: 63.China: Fujian, Guizhou, Hainan (Zhi et al. 2019).(Tsaur & Hsu, 1991)E5961B02-F107-507A-8A23-8CE454C1DB4FTsaur & Hsu in Tsaur et al., 1991a: 21.|China: Taiwan (Tsaur et al. 1991a).(Zhang & Chen, 2011)67C9833E-7F1F-56CF-B2F4-C0367692FBABZhang & Chen, 2011a: 64.|China: Guizhou (Zhang and Chen 2011a).Emeljanov, 20022A3FAE39-9A21-5D85-9A15-2BAFE85FA909Distant, 1907375B3D3B-998D-5604-9709-844CFEC72F28Fennah, 19800EA3DB4A-E064-5804-B9FE-E6E4B7F31DB0Fennah, 1980: 285.China: Hunan; Indonesia: Irian Jaya (Fennah 1980).New record: China: Hunan (Nanyue).Fennah, 1980666BFF28-7E87-5850-953F-FF7AA90A01EAFennah, 1980: 277.China: Hainan; Indonesia: Irian Jaya (Fennah 1980).New record: China: Hainan (Qixianling).Fennah, 1980F090A2EC-A24A-504F-8D6B-DCD39A3E378DFennah, 19805C21B03C-3AF1-5E30-AE09-BE39E0E33A2BFennah, 19801CEF4E28-E1DB-590E-BB73-873D36C862B4Fennah, 1980: 242.China: Hainan; Indonesia: Irian Jaya; Papua New Guinea: Hollandia (Fennah 1980).New record: China: Hainan (Wuzhi Mountain).Walker, 1857AFFFE019-AB44-56D2-9646-F6BFE257086DTsaur & Hsu, 2003F9B5B7B3-CFA3-55DE-B06A-715A4BE0F2D2Tsaur & Hsu, 2003: 438.China: Hainan, Taiwan (Tsaur and Hsu 2003).New record: China: Hainan (Wuzhi Mountain).Tsaur & Hsu, 200371278946-0526-5FDD-97FC-AC27301330C9Tsaur & Hsu, 2003: 436.China: Fujian, Hainan, Taiwan (Tsaur and Hsu 2003).New record: China: Hainan (Liping), Fujian (Meihua).Tsaur & Hsu, 200307061F9B-7191-506E-98CC-48873A2FBD41Tsaur & Hsu, 2003: 438.China: Taiwan (Tsaur and Hsu 2003).(Distant, 1916)449AF2BE-4A78-5E53-A9FA-CEDAF4182C0A,China: Hubei (Fennah 1956); India (Distant 1916).This species is recorded here from China based on female specimens of literature data.Distant, 1906BCA41184-DFC7-5312-AB2A-7E06F895EDC6Zhi & Chen, 2021C9548C7A-58F9-5DBF-879B-336E7471F61EZhi & Chen in Zhi et al., 2021: 7.China: Hunan (Zhi et al. 2021).Zhi & Chen, 2021854B6BD6-ECA4-57DC-9FF0-DC679DC0C083Zhi & Chen in Zhi et al., 2021: 8.China: Yunnan, Guangxi (Zhi et al. 2021).Tsaur & Hsu, 20030B1AF926-9B85-5181-9CD2-5E335AB7203CTsaur & Hsu, 2003: 434.China: Taiwan (Tsaur and Hsu 2003).(Melichar, 1903)D327AD6A-371B-5E51-8F93-D81A71AE0B61Melichar, 1903: 248.|China: Hainan, Taiwan (Tsaur and Hsu 2003); India; Malaysia; Sri Lanka: Peradeniya.New record: China: Hainan (Wuzhi Mountain).Tsaur & Hsu, 20036AB025AB-3F20-5337-A8CA-F3A4D8654124Zhi & Chen, 201766424E8C-DAA6-5CC3-B4D5-01F049F4DE27Zhi & Chen in Zhi et al., 2017: 23.China: Yunnan (Zhi et al. 2017).Zhang & Chen, 20138C795E2B-10E6-5F7D-9DCD-03DF6717D698Zhang & Chen, 2013b: 43.China: Guizhou (Zhang and Chen 2013b).Zhang & Chen, 201372A1DA6E-2AC6-5C85-9606-A8C0144C54B5Zhang & Chen, 2013b: 45.| Zhi et al., 2017: 27.China: Guizhou, Hubei (Zhang and Chen 2013b).Tsaur & Hsu, 2003F8C7F407-C432-5D45-B21A-62EB85C1C8A9Tsaur & Hsu, 2003: 441.China: Zhejiang, Taiwan (Tsaur and Hsu 2003).New record: China: Zhejiang (Fengyang Mountain).Zhi & Chen, 2017393C1576-5A2D-502F-809F-F306AE8FF485Zhi & Chen in Zhi et al., 2017: 30.China: Yunnan (Zhi et al. 2017).Muir, 1922ECDC80C1-C224-5B55-A086-ABA848F27A2AFowler, 19041DA44414-3DA0-584C-B3F9-E496EDCE92A7Ball, 1933951AB21E-6356-503C-B8CA-964D897F7FDDBall, 1933: 473.|China: Guizhou; USA: Delaware, Georgia, Illinois, Iowa, Kansas, Maryland, Massachusetts, Minnesota, Missouri, Nebraska, New Jersey, Oklahoma, South Dakota, Virginia, Wisconsin (Bartlett et al. 2014).Distant, 1906F516293B-AB9D-5F10-8690-4F9D81B21CB3Matsumura, 191438AA8E5E-80E6-5A7E-B0BE-B3499FE66013Matsumura, 1914: 430.| Tsaur et al., 1991a: 76.China: Taiwan (Tsaur et al. 1991a).Stål, 18628BFFA4B0-4570-5363-82F2-6BC9A97EB053Matsumura, 194016C7BBAA-6755-5128-929B-D57F238DAD71Matsumura, 1940: 45.| Tsaur et al., 1991a: 74.China: Taiwan (Tsaur et al. 1991a).Emeljanov, 197143228163-1349-5B53-9275-3A7159115794Hoch, 2006DC7D75CD-D6DA-502A-A302-B82A500118CC(Matsumura, 1911)A9DC88F3-2DF3-54E2-AD1E-ED95EE5554AEMatsumura, 1911: 134.| Van Stalle, 1991: 34.|China: Taiwan (Van Stalle 1991).Emeljanov, 19710A15774F-4A72-586F-9261-EBA8354EC7B6Emeljanov, 2007D90C93E3-6028-5FAA-8D6D-574DBDAE7D48(Matsumura, 1914)6FC9E43C-C7FB-5686-912C-3864478B3584Matsumura, 1914: 423.| Fennah, 1956: 83; Van Stalle, 1991: 46.|China: Taiwan (Matsumura 1914).(Matsumura, 1914)F02EDA15-7BC1-5235-A6FE-063D9578F20AMatsumura, 1914: 425.|China: Taiwan (Matsumura 1914).Emeljanov, 20079B34F8EC-CFE0-52E5-9C13-27A3407A6273(Matsumura, 1914)DE0DF89B-7483-5166-B984-40C68D9B9756Matsumura, 1914: 418.| Schumacher, 1915: 131.| Van Stalle, 1991: 84.| Liang, 2005: 429.|China: Taiwan (Matsumura 1914).(Tsaur, 1990)DFE1B721-7C54-5D8C-B3BB-CBBD3F14FEC9Tsaur, 1990b: 135.|China: Taiwan (Tsaur 1990b).(Fennah, 1956)3BFB83F3-7205-5FBD-9A7C-5E129C08702EFennah, 1956: 451.| Liang, 2005a: 429.|China: Fujian, Guangxi, Hubei (Fennah 1956).New record: China: Guangxi (Baiyangsi).(Tsaur, 1990)FC8BF61C-9D3C-50D4-98FD-1B404BABFFB5Tsaur, 1990b: 137.|China: Taiwan (Tsaur 1990b).(Tsaur, 1989)490B69BF-73DD-5472-AA66-4AAC9429CEBBTsaur, 1989a: 171.| Van Stalle, 1991: 84.|China: Taiwan (Tsaur 1989a).Emeljanov, 20015F512EC6-101E-5A59-980D-745D6C39C418Luo, Liu & Feng 2019BECDBE51-F693-55AA-89C0-8FEBA180E387Luo, Liu & Feng, 2019b: 184.China: Shaanxi (Luo et al. 2019b).Zhi & Chen, 2020ABC6A230-B806-5CF5-B596-0CEF0D09A040Zhi & Chen in Zhi et al., 2020b: 22.China: Yunnan (Zhi et al. 2020b).Guo & Feng, 20108C225D8A-5EBD-5AB6-9B41-12DA2EDF330DGuo & Feng, 2010: 35.China: Gansu (Guo and Feng 2010).Luo, Liu & Feng, 20196B12023A-D946-596F-B2F5-F9A22D142DDBLuo, Liu & Feng, 2019b: 189.China: Guangxi (Luo et al. 2019b).(Distant, 1911)C57FB780-26F4-5717-B931-CF90F3FC2AF6Distant, 1911: 737.| Fennah, 1956: 451.| Van Stalle, 1991: 51.|China: Hubei (Fennah 1956), Guangxi, Hunan, Yunnan (Luo et al. 2019b), Tibet (Guo and Feng 2010); India: Darjeeling (Van Stalle 1991).Zhi & Chen, 20209EF4334D-1F94-5A49-BF0F-E28406861B25Zhi & Chen in Zhi et al., 2020b: 25.China: Yunnan (Zhi et al. 2020b).(Matsumura, 1914)93D7B569-FB05-5852-8017-6A76C2867264Matsumura, 1914: 424.| Tsaur, 1988: 46.| Van Stalle, 1991: 51.|China: Hainan, Taiwan (Guo and Feng 2010).New record: China: Hainan (Diaoluo Mountain).Fennah, 19456B0A7940-7BA2-5AF9-8DE5-B31945E333C0(Mead & Kramer, 1981)872E87FC-4E42-55A4-8500-3FDD55F99E0FMead & Kramer, 1982: 381.|China: Taiwan; Japan; USA: California, New Mexico (Bartlett et al. 2014).This species is recorded here from China based on female specimens of literature data.(Walker, 1851)3FF896B8-0904-5CEA-9754-9FAAE0AE0933Walker, 1851: 343.|China: Hunan; USA: Florida, Colorado; Georgia; Illinois; Maryland; Massachusetts; Mississippi; New Jersey; North Carolina; South Carolina; Texa (Bartlett et al. 2014).This species is recorded here from China based on female specimens of literature data.Emeljanov, 1971372F7A3B-9248-5955-91EB-15A82C524580Van Stalle, 19913837287A-9DA6-5545-A88B-E975D9FF9122Van Stalle, 1991: 22.| Guo et al., 2009: 48.China: Hainan, Henan, Sichuan (Guo et al. 2009), Guizhou; Malaysia: Borneo, Pahang (Van Stalle 1991).New record: China: Guizhou (Duyun). This species is recorded here from China only based on female specimens.(Tsaur, Hsu & Van Stalle, 1988)714BFD67-FCB0-5DCB-8D69-E615B4F8E8C6Tsaur, Hsu & Van Stalle, 1988: 52.|China: Fujian, Taiwan (Tsaur et al. 1988).New record: China: Fujjian (Shaowu).(Tsaur, Hsu & Van Stalle, 1988)A646C45B-654E-50C8-99D9-1919D68F09FCTsaur, Hsu & Van Stalle, 1988: 50.|China: Fujian, Taiwan (Tsaur et al. 1988).New record: China: Fujjian (Shaowu).(Tsaur, Hsu & Van Stalle, 1988)DE2C66BA-BF7B-52EF-8784-7CD89FC3A020Tsaur, Hsu & Van Stalle, 1988: 53.|China: Guangxi, Taiwan (Tsaur et al. 1988); Japan: Honshu (Hayashi and Fujinuma 2016).New record: China: Guangxi (Lingchuan).Zhi & Chen, 20186644832B-95D5-51EC-8EBA-2DC149443E2CZhi & Chen in Zhi et al., 2018a: 5.China: Yunnan (Zhi et al. 2018a).Matsumura, 19144837A9C9-CCA8-5DE7-8E41-1D7BE8F2855BMatsumura, 1914: 426.| Van Stalle, 1991: 23.| Guo et al., 2009: 50.| Zhi et al., 2018a: 9.China: Guangxi, Yunnan (Zhi et al. 2018a), Taiwan (Van Stalle 1991).(Noualhier, 1896)C6F3EA86-F66C-5F35-B4A7-237610233A31Noualhier, 1896: 255.| Fennah, 1956: 455.|China: Hainan, Sichuan, Yunnan (Guo et al. 2009); Cambodia: (Noualhier 1896).New record: China: Yunnan (Yaoqu, Mengla, Longling, Kunming), Sichuan (Yaan), Hainan (Yinggeling).Zhi & Chen, 2018BF059895-42F7-5A75-A592-AD29073A89B7Zhi & Chen in Zhi et al., 2018a: 9.China: Yunnan (Zhi et al. 2018a).(Jacobi, 1944)CC5D8363-DCC2-5B5F-9C73-831542D62556Jacobi, 1944: 12.| Chou, 1985: 23.|China: Beijing, Anhui, Fujian (Jacobi 1944), Guangdong, Guangxi (Guo et al. 2009), Henan, Hubei, Hunan, Sichuan, Guizhou, Zhejiang, Taiwan; Cambodia; Japan: Kyushu (Hayashi and Fujinuma 2016).New record: China: Beijing (Mentougou), Hunan (Chenzhou, Huping), Fujian (Fuzhou), Guangxi (Lingchuan), Guangdong (Lohchan).Guo & Wang, 2009622FEAE2-33D8-5D55-8FA4-D4B576743D34Guo & Wang in Guo et al., 2009: 54.China: Shaanxi (Guo et al. 2009).Guo & Wang, 20098ED00389-A5E0-5283-8807-61F71C258082Guo & Wang in Guo et al., 2009: 54.China: Shaanxi (Guo et al. 2009), Gansu.New record: China: Shaanxi (Hanzhong), Gansu (Xiaolong Mountain).Guo & Wang, 2009A6ABBBED-2B4B-5EF6-92FA-FD44E56AFF81Guo & Wang in Guo et al., 2009: 56.China: Fujian, Shaanxi, Henan, Hunan (Guo et al. 2009), Yunnan.New record: China: Yunnan (Lvchun).(Ishihara, 1961)650F8AC3-4E94-5444-8D65-A768BFB7A164Ishihara, 1961: 228.|China: Guizhou (Zhi et al. 2018a), Yunnan; Thailand: Doi Inthanon (Van Stalle 1991).New record: China: Yunnan (Sumie).(Noualhier, 1896) comb. nov.9EB60169-8829-5994-B4C1-DCAA23ECF8A2Noualhier, 1896: 255.| Jacobi, 1917: 11.| Fennah, 1956: 453.|China: Guangdong, Hubei (Fennah 1956); Cambodia (Van Stalle 1991).This species was originally belonged to , and when the authors observed the paratype specimens of this species, we found that its morphology indicates the misclassification of this species, this species with strongly elevated and foliaceous lateral carinae, consistent with the diagnostic characteristics of , so in this study this species was transferred to as a new combination.Stål, 1862DBB9B0D5-FB48-5A5E-99BA-124908EBC624Kato, 1932A0D67427-A39E-5EDC-A723-9788E1C22167Kato, 1932: 216.China: Northwestern of China (Kato 1932).(Distant, 1911)6DD142DE-48F5-5F91-BC0F-97CB3FAB9F78Distant, 1911: 738.|China: Yunnan; Sri Lanka: Trincomalee (Distant 1911); USA: Puerto Rico.New record: China: Yunnan (Yuanmou).Distant, 1911E82674FB-FEFC-587D-A5B9-4793DCD9B350Distant, 1911: 735.| Van Stalle, 1991: 80.China: Beijing; India: (Van Stalle 1991).New record: China: Beijing (Xishan).Van Stalle, 198760E32BE8-7062-5E1C-B751-9C3F1C74D968Van Stalle, 1987: 66.China: Guangxi, Hubei; Malawi; Tanzania; Zimbabwe: ex Rhodesia (Van Stalle 1987).New record: China: Guangxi (Longsheng), Hubei (Shennongjia).Matsumura, 19148B86B655-1939-5F8C-8051-0BF8D984A1BEMatsumura, 1914: 424.China: Taiwan (Matsumura 1914).Wang, 1991DCCAD982-D5BC-5594-B2BE-DD07C6A07D2CWang, 1991: 85.China: Hebei (Wang 1991).Emeljanov, 2001F0C80414-8BE8-5C6E-A80B-E117D88E93FB(Van Stalle, 1991)F9E92CB3-0565-5575-A2F5-F4CC4BA6BADBVan Stalle, 1991: 72.|China: Tibet; Sri Lanka: (Van Stalle 1991).Kirschbaum, 1868CB5C33E1-8FA5-51B9-9B4A-AAF5961A5EB9(Uhler, 1896)8F546EFF-29BA-524A-B1A2-D8575630C03AUhler, 1896: 281.|China: Beijing (Liang 2005b), Shanghai, Fujian, Jiangxi, Jiangsu, Shaanxi, Sichuan, Zhejiang; Japan: Hokkaido, Honshu, Kyushu, Shikoku (Hayashi and Fujinuma 2016); Russia: Khabarovsk, Primorye.New record: China: Hebei (Shijiazhuang), Jiangsu (Jinshan), Shanghai (Songjiang).(Stål, 1859)76A00D13-EC3F-53E9-9C38-FC4F58348173Stål, 1859: 272.|China: Hainan, Hongkong (Van Stalle 1991).New record: China: Hainan (Xisha).(Linné, 1761)5AE3C81D-91CF-5BA8-801C-35B67142779FLinné, 1761: 242.|China: Nei-Mongol, Heilongjiang; Iran: bādeh, Albāji, Bampur, Bazmān, Birjand, Chābahār, Dālaki, Evin, Gambuyeh, Gāvbandi, Gharechaman, Hafttappeh, Hamidieh, Hāresābād, Hashtpar, Irānshahr, Kandovān (Māzandarān), Marand, Miāneh-ZanjānRd, Minushahr, Mollāsāni, Shādegān, Shieh, Susangerd, Suza, Tabriz, Varāmin, Zābol (Kalkandelen 1990); Afghanistan (Nast 1972, Holzinger et al. 2003); Albania; Algeria: (Nast 1972, Holzinger et al. 2003) Armenia; Austria; Azerbaijan (Nast 1972, Holzinger et al. 2003); Belgium: (Nast 1972, Holzinger et al. 2003); Cyprus; Czech Republic; Denmark: (Nast 1972, Holzinger et al. 2003); Estonia: (Nast 1972, Holzinger et al. 2003); Finland: (Nast 1972, Holzinger et al. 2003); France: (Nast 1972, Holzinger et al. 2003); Georgia: (Nast 1972, Holzinger et al. 2003); Germany: (Nast 1972, Holzinger et al. 2003); UK: (Nast 1972, Holzinger et al. 2003); Greece: (Nast 1972, Holzinger et al. 2003); Hungary: (Nast 1972, Holzinger et al. 2003); Ireland: (Nast 1972, Holzinger et al. 2003); Israel: (Nast 1972, Holzinger et al. 2003); Italy: (Nast 1972, Holzinger et al. 2003); Jordan: (Nast 1972); Kazakhstan: (Holzinger et al. 2003); Kyrgyzstan: (Nast 1972, Holzinger et al. 2003); Libya: (Nast 1972); Lithuania; Moldova: (Holzinger et al. 2003); Mongolia; Netherlands: (Nast 1972, Holzinger et al. 2003); Poland: (Nast 1972, Holzinger et al. 2003); Portugal: (Nast 1972, Holzinger et al. 2003); Romania: (Nast 1972, Holzinger et al. 2003); Russia: Primorye; Slovakia; Spain; Sweden; Switzerland; Tadzhikistan: (Nast 1972, Holzinger et al. 2003); Tunisia (Nast 1972); Turkey (Kalkandelen 1990); Turkmenistan (Kalkandelen 1990); Ukraine: (Nast 1972, Holzinger et al. 2003); Yugoslavia:(Nast 1972, Holzinger et al. 2003).(Matsumura, 1914)048A3ED1-DB16-5E75-831A-96563D02A93DMatsumura, 1914: 420.| Schumacher, 1915: 131.|China: Hainan, Taiwan (Van Stalle 1991); Nansei-shoto: Ryukyu Islands (Hayashi and Fujinuma 2016).New record: China: Hainan (Lingshui).(Muir, 1925)5FF2ECDC-4A01-5509-A6C1-38A4C3CE5940Muir, 1925: 365.|China: Jiangsu, Hubei (Fennah 1956); Japan (Fennah 1956).Emeljanov, 197114B256F2-310B-5756-8933-08AE993BB69B(Dlabola, 1985)F38B23E2-F600-55B7-9500-A5CD0477D3BEDlabola, 1965: 87.|China: Beijing; Mongolia: Uburchangaj aimak (Anonymous 2015).New record: China: Beijing (Mentougou).Guo & Wang, 20070B88D95B-EDDA-5457-8ED2-119DDABFA6FEGuo & Wang, 2007: 276.| Bai et al., 2015: 37.China: Fujian (Bai et al. 2015), Hubei, Hunan (Guo and Wang 2007), Zhejiang, Sichuan, Guizhou, Hebei, Qinghai, Jiangsu.New record: China: Jiangsu (Xinghua), Jiangsu (Zhenze).(Matsumura, 1914)2717FFE0-6475-5B44-9B4F-4AFDFC4B9B7FMatsumura, 1914: 419.|China: Guizhou, Hunan; South Korea: Gyeongsangubuk-do (Rahmam et al. 2011); Japan: Honshu, Kyushu (Hayashi and Fujinuma 2016).New record: China: Guizhou (Duyun), Guizhou (Tongren), Guizhou (Kaili), Hunan (Suining).(Matsumura, 1914)9C995D48-7FCE-51A7-874D-9D86083F13DBMatsumura, 1914: 419.| Chou, 1985: 20.|China: Beijing, Anhui, Fujian, Hunan, Hubei, Jinlin, Shaanxi, Zhejiang, Jiangsu, Shanghai, Sichuan, Guizhou; Japan: Honshu, Kyushu, Shikoku (Hayashi and Fujinuma 2016); Russia: Primorye; South Korea: Daegu (Rahmam et al. 2011).New record: China: Anhui (Anhui labor university), Shaanxi (Foping nature reserve, Taibai Mountain, Ningqiang, Shiquan, Suining, Chenxi), Hunan (Hupengshan nature reserve, Zhangjiajie nature reserve), Hubei (Houhe nature reserve, Shennongjia), Jilin (Linjiang), Fujian (Shaowu, Huangkeng, Jianning, Daan), Anhui (Anhui labor university), Zhejiang (Fengyangshan), Beijing (Mentougou), Zhejiang (Hangzhou), Jiangsu (Zhenze, Suzhou), Shanghai (Qingpu, Bao Mountain, Sheshan, Jinshan), Sichuan (Qianjiang).(Dufour, 1833)C8C3CC1B-1C5B-5AA1-B3C8-1264EA18C1BDDufour, 1833: 224.|China: Chongqing; Armenia; Austria; Bulgaria; Czech Republic; France; Germany; Greece; Hungary: Andornaktalya; Iran: North (Mozaffarian and Wilson 2011); Italy: Emilia Romagna, Piemonte; Kazakhstan; Portugal; Romania: Csı ´kszereda, Fundulea; Russia: Azov; Serbia: Vršac, Topla, Rajac, South Banat District; Slovakia; Spain; Tadzhikistan; Turkey; Ukraine; Yugoslavia.New record: China: Chongqing.Bai, Guo & Feng, 20152C54E1A3-8C56-569D-9ECF-3064DF619F1CBai, Guo & Feng, 2015: 38.China: Anhui, Sichuan, Zhejiang (Bai et al. 2015).Emeljanov, 20077C738C72-1D63-576A-A240-37DCCE0A647D(Matsumura, 1914)A0B4BD3E-6724-538C-BE0E-DE4821EFB11CMatsumura, 1914: 427.| Van Stalle, 1991: 31.| Schumacher, 1915: 131.|China: Taiwan (Matsumura 1914).(Jacobi, 1944)0DDF7FC0-CB0E-5549-8000-870C3C256372Jacobi, 1944: 13.| Fennah, 1956: 454.|China: Fujian (Jacobi 1944), Guangdong, Hongkong, Hubei, Sichuan (Fennah 1956), Yunnan, Tibet, Guangxi, Guizhou, Hunan, Taiwan.(Tsaur, Hsu & Van Stalle, 1988)9383C656-69FC-50C4-AE0C-56A55E9310B3Tsaur, Hsu & Van Stalle, 1988: 41.| Van Stalle, 1988: 29.|China: Taiwan (Tsaur et al. 1988).Emeljanov, 2002FBFC17DE-073E-55D6-8618-E563C0552F16Matsumura, 1914B562C45F-0AC5-505A-9DD5-FEC7D0AD617AZhang & Chen, 20113ECEAD8C-6927-5C0D-87E5-2E2B5176C8BDZhang & Chen, 2011b: 53.China: Sichuan (Zhi et al. 2020a), Guangxi, Guizhou (Zhang and Chen 2011b), Xinjiang, Yunnan (Zhi et al. 2020a).New record: China: Xinjiang (Changji Temple), Guangxi (Lintian).Matsumura, 1914371E1BC6-F487-5B61-AC3B-AAE7FD621361Matsumura, 1914: 417.| Hori, 1982: 181.| Tsaur et al., 1991b: 37.| Zhang & Chen, 2011b: 50.| Hayashi & Fujinuma, 2016: 323.China: Fujian, Zhejiang, Taiwan (Zhang and Chen 2011b); Japan; Nansei-shoto: Ryukyu Islands (Hayashi and Fujinuma 2016).New record: China: Fujian (Taoyuan valley scenic spot of wuyi Mountain).Matsumura, 1914EC017CC6-4405-5320-BC10-7634F0C17C7DMatsumura, 1914: 415.| Hori, 1982: 181.| Tsaur et al., 1991b: 39.China: Zhejiang, Taiwan (Matsumura 1914).New record: China: Zhejiang (Jiulong Mountain, Wuyanling).(Matsumura, 1914)7A97A4B6-E069-5308-843C-3CAF9ADB3C5DMatsumura, 1914: 416.China: Taiwan (Matsumura 1914).Tsaur & Hsu, 1991B6D64390-B163-5DCF-B070-0ECFF462B375Tsaur & Hsu in Tsaur et al., 1991a: 29.China: Shaanxi, Zhejiang, Yunnan, Taiwan (Tsaur et al. 1991a).New record: China: Zhejiang (Fengyang Mountain).Fennah, 1956387E6F11-676E-5101-B53B-A7246A3EF440Fennah, 1956: 458; Zhang & Chen, 2011b: 50.China: Guangdong (Fennah 1956).Zhang & Chen, 2011B0735B12-8826-5BB9-AC0D-C1FA08E668EDZhang & Chen, 2011b: 54.China: Guizhou (Zhang and Chen 2011b).Hori, 198269889B0C-1100-54D2-9CD0-8656D17437F1Hori, 1982: 179.| Tsaur et al., 1991a: 41.| Zhang & Chen, 2011b: 50.China: Zhejiang, Taiwan (Tsaur et al. 1991a).New record: China: Zhejiang (Fengyang).Tsaur & Hsu, 19917B432300-AA91-5F22-8411-20CFB1D98202Tsaur & Hsu in Tsaur et al., 1991a: 44.| Zhang & Chen, 2011b: 50.China: Fujian, Taiwan (Tsaur et al. 1991a).New record: China: Fujian (Longyan City Contour Park).Tsaur & Hsu, 1991B11F5E96-C220-58F4-A9B7-91BFF799A547Tsaur & Hsu in Tsaur et al., 1991a: 28.China: Yunnan, Taiwan (Tsaur et al. 1991a).New record: China: Yunnan (Yangyang Valley, Matang Reservoir).Zhi, Zhang, Yang & Chen, 20206FA31B19-DE44-5A14-945E-F7A1591121EFZhi, Zhang, Yang & Chen, 2020a: 8.China: Yunnan (Zhi et al. 2020a).Tsaur & Hsu, 1991BC2FDE65-2DC5-5F98-80A3-F6A484D75011Tsaur & Hsu in Tsaur et al., 1991a: 31.China: Fujian, Sichuan, Taiwan (Tsaur et al. 1991a).New record: China: Fujian (Wuyi Mountain), Sichuan (Emei Mountain).Zhi, Zhang, Yang & Chen, 202036145D78-AD41-5AB9-A980-89D2D1360AD6Zhi, Zhang, Yang & Chen, 2020a: 11.China: Yunnan (Zhi et al. 2020a).Hori, 198233F280E6-1E48-510F-8254-EEA650569DE8Hori, 1982: 176.| Tsaur et al., 1991a: 35.| Zhang & Chen, 2011b: 50.China: Yunnan, Taiwan (Tsaur et al. 1991a).New record: China: Yunnan (Yangyang Valley, Matang Reservoir).Fennah, 1956DB1C8EDB-3BCA-58A9-B5BD-96327C975828Fennah, 1956: 458.China: Zhejiang (Fennah 1956).Fennah, 1956F6669645-CBC6-590D-8098-25E563C46C0BFennah, 1956: 457.China: Guangdong (Fennah 1956).Fennah, 1956F62DA16A-AFA8-53BF-858B-8024AD3B8B1CFennah, 1956: 457.China: Hubei (Fennah 1956).Matsumura,1914BBD47E53-5BB4-554F-B004-1FF59C75C034Matsumura, 1914: 414.| Chou, 1985: 23; Chou, 1998: 382.| Liang, 2005b: 429.| Zhang & Chen, 2011b: 50.|China: Fujjian, Guizhou, Guangxi, Guangdong, Hainan, Hunan, Sichuan (Liang 2005b), Yunnan, Zhejiang; Japan: Honshu, Kyushu, Shikoku (Hayashi and Fujinuma 2016).New record: China: Hainan (Diaoluoshan, Limu Island), Fujian (Wuyi Mountain, Chongan, Guangze, Yongan), Sichuan (Emei Mountain), Guangxi (Huaping nature reserve), Hunan (Shenzhou); Guangdong (Dinghu Mountain), Zhejiang (Qingyuan, Longquan).Matsumura, 1914BA74F6E8-0C47-55E1-B58D-22507F108C95Matsumura, 1914: 412.| Esaki 1932: 1774.| Hori, 1982: 181.| Tsaur et al., 1991b: 33.China: Yunnan, Fujian, Taiwan (Tsaur et al. 1991a).New record: China: Fujian (Shaowu Jiangshi Nature Reserve).Jacobi, 1944F74FBC5A-7A57-5448-82F5-15BBB71C094CJacobi, 1944: 15.| Fennah, 1978: 213.China: Fujian (Jacobi 1944); Vietnam: Hanio (Fennah 1978).Matsumura, 1914FD96DF70-28BA-5890-A052-633254C51AA7Matsumura, 1914: 417.|China: Guangdong, Taiwan (Tsaur et al. 1991a).New record: China: Guangdong (Shaoguan Nanling).Jacobi, 1944FB8B66DF-2902-5884-AEB8-6F0BE4235395Jacobi, 1944: 15.| Zhang & Chen, 2011b: 50.China: Fujian (Jacobi 1944).Hori, 19828F93EDED-0633-51FF-AC3F-F225587812C0Hori, 1982: 178.| Tsaur et al., 1991a: 48.| Zhang & Chen, 2011b: 50.China: Yunnan, Taiwan (Tsaur et al. 1991a).New record: China: Yunnan (Mengla).Tsaur & Hsu, 1991884E2AE0-57E2-5A42-AAE9-B4A4120AEF6ETsaur & Hsu in Tsaur et al., 1991a: 46.| Zhang & Chen, 2011b: 50.China: Fujian, Hainan, Taiwan (Tsaur et al. 1991a).New record: China: Fujian (Longyan), Hainan (Jianfengling).Jacobi, 1944583020E7-D2B4-5AB0-9BE3-08F8347C9AAFJacobi, 1944: 14| Zhang & Chen, 2011b: 50.China: Fujian (Jacobi 1944).Distant, 1906FB87F526-3FC0-5A82-B43F-DCC310C70806Tsaur & Hsu, 199165535261-09A3-56A8-9C5B-BDD54C5EEF38Tsaur & Hsu in Tsaur et al., 1991a: 59.China: Fujian, Taiwan (Tsaur et al. 1991a).New record: China: Fujian (Fengyang Mountain).(Matsumura, 1900)129E9CF7-62D5-5ABA-BB45-23871731E7D1Matsumura, 1900: 208.|China: Gansu, Jilin; Japan: Hokkaido, Honshu, Kyushu, Shikoku (Hayashi and Fujinuma 2016); Korea; Russia: Sakhalin.New record: China: Gansu (Wen County).Tsaur & Hsu, 19919C958D3C-7652-57F4-8D56-BB710E37238ATsaur & Hsu in Tsaur et al., 1991b: 61.China: Jilin, Fujian, Hunan, Taiwan (Tsaur et al. 1991a).New record: China: Fujian (Wuyi Mountain), Hunan (Huping Mountain).Luo, Liu & Feng, 201977B700A1-797F-581D-B651-806F8D1B5297Luo, Liu & Feng, 2019a: 140.China: Shaanxi (Luo et al. 2019a), Henan, Gansu.Anufriev, 19874E89264D-560C-5DE2-8899-690247C9AA5EAnufriev, 1987: 15.| Anufriev & Emeljanov, 1988: 449.China: Jilin, Fujian, Hunan, Tibet, Taiwan; Russia: Kuril Islands (Anufriev 1987).New record: China: Tibet (Motlin Green).(Metcalf, 1936)6A6A657F-5A45-537B-AD4C-4DD814A2C982Melichar, 1902: 85.|China: Sichuan (Anufriev 1987).Matsumura, 1914C4A02E79-A942-5714-A059-FF2FCE4F1BE4Matsumura, 1914: 411.China: Taiwan (Matsumura 1914).Luo, Liu & Feng, 2019EAE4AA75-930F-5AE3-B29D-753E8A91C00ALuo, Liu & Feng, 2019a: 144.China: Zhejiang (Luo et al. 2019a).Matsumura, 19141BB998C4-309B-5675-A196-2FDF0E36E6B9Matsumura, 1914: 409.|China: Jilin, Guangxi, Shaanxi, Sichuan; Japan: Hokkaido, Honshu (Hayashi and Fujinuma 2016); Russia: Far East (Anufriev 1987).New record: China: Jilin (Antu County), Guangxi (Longsheng), Sichuan (Yaan).Distant, 190696AA91AF-3556-5F60-8835-FBEEF4AEF377Distant, 1906: 261.| Anufriev, 1987: 6.China: Liaoning, Shaanxi; Myanmar: (Distant 1906).New record: China: Liaoning (Baling County National Balding National Nature Reserve), Shaanxi (Shiquan, Qinling).Tsaur & Hsu, 1991F45EE93D-1970-5AB4-843B-E3D32669EACCTsaur & Hsu in Tsaur et al., 1991a: 55.China: Beijing, Fujian, Taiwan (Tsaur et al. 1991a).New record: China: Beijing (Mentougou), Fujian (Meihua).Tsaur & Hsu, 19916EEDB20E-7200-5845-8AEF-7DB1C47D2D5BTsaur & Hsu in Tsaur et al., 1991a: 50.China: Hainan, Shaanxi, Yunnan, Ningxia, Zhejiang, Tibet, Taiwan (Tsaur et al. 1991a).New record: China: Zhejiang (Fengyang Mountain, Linan), Hainan (Yinggeling), Tibet (Yadong); Shaanxi (Huayin), Yunnan (Lvchun), Ningxia (Liupan Mountain).Matsumura, 191491C92FDC-C07B-5578-BF0A-AD874DABE762Matsumura, 1914: 410.China: Hubei, Jiangxi, Hunan, Jilin, Taiwan (Tsaur et al. 1991a).New record: China: Hubei (Shennongjia), Jiangxi (Jinggang Mountain), Hunan (Mang Mountain), Hubei (Shennongjia), Jilin (Changbai Mountain).Matsumura, 1914FF492551-9878-5301-8E1C-6CA89A5E770EMatsumura, 1914: 410.| Anufriev, 1987: 18.China: Yunnan, Jiangxi, Zhejiang, Hunan, Tibet, Taiwan (Matsumura 1914).New record: China: Hunan (Qianyang), Jiangxi (Lu Mountain), Yunnan (Mengla Longmen, Matang Reservoir), Zhejiang (Hangzhou), Tibet (Langxian Cuona).Tsaur & Hsu, 1991251304F6-F611-5954-AC49-EF09A0D16160Tsaur & Hsu in Tsaur et al., 1991a: 57.China: Yunnan, Taiwan (Tsaur et al. 1991a).New record: China: Yunnan (Tengchong Laifeng Mountain).(Vilbaste, 1968)3602C65F-A0E0-58BA-80DF-D9921DEC7E14Vilbaste, 1968: 9.|China: Sichuan; Russia: Primorsky Territory, South of the Khabarovsk Territory (Anufriev 1987); Japan: Hokkaido.Anufriev, 1987FFA1C0F2-3824-5C85-A022-9DD879306BB7Anufriev, 1987: 7.| Anufriev & Emeljanov, 1988: 448.| Anufriev, 2009: 68.China: Shaanxi, Zhejiang, Tibet; Russia: Primorye (Anufriev 1987).New record: China: Tibet (Bomi Yigong, Yadong), Shaanxi (Hua Mountain, Huxian), Zhejiang (Tianmu Mountain).Rahman, Kwon & Suh, 2017CA31306D-26BD-5B62-BFA3-2B84C19DB6A4Rahman, Kwon & Suh, 2017: 10.China: Guizhou; South Korea: Gyeongsangbuk-do (Rahman et al. 2017).New record: China: Guizhou (Qiandong).Metcalf, 19383FEB49C8-1220-5935-B0E4-2B3DBE3A1E45Muir, 19220CB5516F-E7C6-566A-8782-904DD3844A0ATsaur, 19892E9092F0-220C-584F-8FE8-25C805F4E67DTsaur, 1989: 82.China: Taiwan (Tsaur 1989b).
Analysis
Checklist
Ten cixiid tribes are reported in China: Spinola, 1839, Muir, 1922, Metcalf, 1938, Metcalf, 1938, Emeljanov, 1971, Emeljanov, 1989, Emeljanov, 2002; Emeljanov, 2002, Emeljanov, 2002, and Emeljanov, 2002. These tribes include 35 genera and subgenera, 250 species and 400 collection records from 28 Chinese provinces. In this study, 77 new species were recorded for the first time from China.
Regional richness and endemism
A species richness gradient occurs from north to south and from west to east for as shown in Fig. 2. Substantial variation in species richness and endemism among the different zoogeographic regions were observed. Table 1 describes the species richness of by region, ranging from 5 species and no endemic species in the Nei Mongol-Xinjiang region, to 161 species and 69.57% of endemic species in Taiwan region. In-between, species richness and endemism ratios are distributed in two groups: the Northeast China and the Qinghai-Tibet regions, respectively with 8 and 10 species and 12.5% and 20% of endemism, and the North, Southwest and Central China regions, which have comparable numbers of species and endemism, respectively, with 29, 43 and 60 species and 33-40% of endemism. No significant differences in endemism among regions was observed. More than five-fourths of the species (205 species; 82%) are reported to occur in only in China, depicting a high level of endemism of the Chinese fauna for this family (Table 1).
Figure 2.
The distribution of species records of species in China.
Table 1.
Species richness, endemism and the proportion of endemic species in the 8 zoogeographical regions of China.
Zoogeographical regions
Species richness
Number of endemic species
Endemic species%
South China
78
14
17.95
Southwest China
43
17
39.53
Central China
60
20
33.33
North China
29
10
34.48
Northeast China
8
1
12.5
Nei Mongol-Xinjiang
5
0
0
Qinghai-Tibet
10
2
20.00
Taiwan
161
112
69.57
China
250
205
82.00
Distribution patterns of cixiid species in China
Based on the eight zoogeographic regions of China (Fig. 1), 38 main distribution patterns are observed (Table 2). The number of species distributed in a single region (accounting for regional endemism) is highly variable among the regions: Taiwan (44.80%), Central China (8.00%), Southwest China (6.80%), South China (5.60%), North China (4.00%), Qinghai-Tibet (0.80%) and Northeast China (0.40%). No endemic species were observed in the Nei Mongol-Xinjiang region (Table 2). Nine bi-regional distribution patterns were observed, and among them the South China-Taiwan pattern has the greatest number of species (15 species, 6.00% of the species). Nine tri-regional distribution patterns were also observed, among which, the largest number of species (11 species, 4.40% of the species) was for the Central-South China-Taiwan distribution pattern. The Southwest-South China-Taiwan distribution pattern is depicted by 6 species (2.40% of the species). Five distribution patterns occur in 4 zoogeographic regions, among which the North Southwest-Central-South China region and the Northeast-Central-South China-Taiwan region have two species (0.80% of the total number of cixiids in China). All the remaining four-, five-, six- and seven-regional distribution patterns have only a single species, accounting for 0.40% of the total number of cixiids in China (Table 2).
Table 2.
Distribution patterns of among China zoogeographical regions and proportion of species in these patterns of the total number of species.
* Abbreviations: NEC, Northeast China; NC, North China; NX, Nei Mongol-Xinjiang; QT, Qinghai-Tibet; SWC, Southwest China; CC, Central China; SC, South China; TW, Taiwan.
Distributed pattern
Number of species
Species number %
TW
112
44.80
CC
20
8.00
SWC
17
6.80
SC
14
5.60
NC
10
4.00
QT
2
0.80
NEC
1
0.40
SC-TW
15
6.00
SWC-SC
3
1.20
SWC-CC
3
1.20
CC-SC
2
0.80
NC-CC
2
0.80
CC-TW
1
0.40
NEC-NC
1
0.40
NC-SC
1
0.40
NEC-NX
1
0.40
CC-SC-TW
11
4.40
SWC-SC-TW
6
2.40
NC-SC-CC
2
0.80
SWC-SC-CC
2
0.80
NC-SC-TW
2
0.80
NX-SC-TW
2
0.80
NC-QT-CC
1
0.40
NX-CC-SC
1
0.40
QT-SC-TW
1
0.40
NC-SWC-CC-SC
2
0.80
NEC-CC-SC-TW
2
0.80
NC-CC-SC-TW
1
0.40
QT-SWC-CC-SC
1
0.40
SWC-CC-SC-TW
1
0.40
NC-SWC-CC-SC-TW
2
0.80
NEC-NC-SWC-CC-SC
1
0.40
NC-QT-SWC-CC-SC
1
0.40
NEC-QT-CC-SC-TW
1
0.40
QT-SWC-CC-SC-TW
1
0.40
NEC-NC-SWC-CC-TW
1
0.40
NC-QT-SWC-CC-SC-TW
1
0.40
NC-NX-QT-SWC-CC-SC-TW
1
0.40
Cixiid patterns of distribution at the tribal level and generic level
Of the ten cixiid tribes distributed in China (Table 3), (21.20%, Fig. 6b) and (17.20%, Fig. 7a) are the two most widely distributed tribes in China. (Fig. 5a), which is the most species-rich tribe with 45.20% of the species, is distributed in 7 regions of China, but has not been reported from the Northeast China region. (5.20%, Fig. 3a) is not distributed in the Palaearctic realm in China; (6.40%, Fig. 5b) and (2.00%, Fig. 4a) are distributed only in the Southwest, Central, South China, and in the Taiwan regions. The remaining tribes, (0.40%, Fig. 3b), (0.80%, Fig. 4b), (1.20%, Fig. 6a) and (0.40%, Fig. 7b) are only found in Taiwan.
Table 3.
Number and percentage of cixiid species distributed in China by tribes among the Chinese zoogeographical regions. Abbreviations: NEC, Northeast China; NC, North China; NX, Nei Mongol-Xinjiang; QT, Qinghai-Tibet; SWC, Southwest China; CC, Central China; SC, South China; TW, Taiwan.
Chinese tribes of Cixiidae
Number of species
Species %
Zoogeographical distribution
Andini Emeljanov, 2002
13
5.20
NC, QT, SWC, CC, SC, TW
Bennini Metcalf, 1938
1
0.40
TW
Brixiini Emeljanov, 2002
2
0.80
TW
Cixiini Spinola, 1839
113
45.20
NC, NX, QT, SWC, CC, SC, TW
Eucarpiini Emeljanov, 2002
16
6.40
SWC, CC, SC, TW
Oecleini Muir, 1922
3
1.20
TW
Pentastirini Emeljanov, 1971
53
21.20
NEC, NC, NX, QT, SWC, CC, SC, TW
Semonini Emeljanov, 2002
43
17.20
NEC, NC, NX, QT, SWC, CC, SC, TW
Stenophlepsiini Metcalf, 1938
1
0.40
TW
Borysthenini Emeljanov, 1989
5
2.00
SWC, CC, SC, TW
Figure 6.
Distribution of the tribes and in China. (a) Muir, 1922; (b) Emeljanov, 1971.
Figure 7.
Distribution of the tribes and in China. (a) Emeljanov, 2002; (b) Metcalf, 1938.
Figure 5.
Distribution of records of the tribes and in China. (a) Spinola, 1839; (b) Emeljanov, 2002.
Figure 3.
Distribution of records of the tribes and in China. (a) Emeljanov, 2002; (b) Metcalf, 1938.
Figure 4.
Distribution of records of the tribes and in China. (a) Emeljanov, 1989; (b) Emeljanov, 2002.
Thirty-three genera are present in China (Table 4; Fig. 3; Fig. 4; Fig. 5; Fig. 6; Fig. 7), with (18 species, 7.2%) being the most widespread genus in China although (5 species, 2%) is only unreported from the Qinghai-Tibet region. is the most diverse (95 species, 38%) but is not distributed in the Southwest and Northeast China regions. (25 species, 10%) is not distributed in the Northeast China and Qinghai-Tibet regions. and (each with 6 species 2.4%) are both undistributed in the Nei Mongol-Xinjiang region, while the former is not reported from the Qinghai-Tibet region and the latter not reported from the Taiwan region. (1 species, 0.4%) is only distributed in the Qinghai-Tibet region and 10 genera are only reported from the Taiwan region. In addition, we also found that 16 genera are all distributed in the south of Sino-Japanese/Oriental boundary.
Table 4.
Number and percentage of cixiid genus and species distributed in China by genera amongst the Chinese zoogeographical regions. Abbreviations: NEC, Northeast China; NC, North China; NX, Nei Mongol-Xinjiang; QT, Qinghai-Tibet; SWC, Southwest China; CC, Central China; SC, South China; TW, Taiwan.
Chinese genera of Cixiidae
Number of species
Species
Zoogeographical distribution
%
Andes Stål, 1866
10
4
NC, QT, SWC, CC, SC, TW
Andixius Emeljanov & Hayashi, 2007
3
1.2
SWC, TW
Ankistrus Tsaur & Hsu, 1991
7
2.8
TW
Arosinus Emeljanov, 2007
2
0.8
TW
Atretus Emeljanov, 2007
5
2
CC, SC, TW
Bajauana Distant, 1907
2
0.8
CC, SC
Betacixius Matsumura, 1914
25
10
NC, NX, SWC, CC, SC, TW
Borysthenes Stål, 1866
5
2
SWC, CC, SC, TW
Brixia Stål, 1859
2
0.8
TW
Cixius Latreille, 1804
95
38
NC, NX, QT, CC, SC, TW
Dilacreon Fennah, 1980
1
0.4
SC
Eucarpia Walker, 1857
4
1.6
SC, TW
Euryphlepsia Muir, 1922
1
0.4
TW
Gonophallus Tsaur & Hsu, 1991
1
0.4
TW
Haplaxius Fowler, 1904
1
0.4
CC
Indolipa Emeljanov, 2001
7
2.8
NC, QT, SWC, CC, SC, TW
Kirbyana Distant, 1906
4
1.6
SWC, CC, SC, TW
Kotonisia Matsumura, 1938
1
0.4
TW
Kuvera Distant, 1906
18
7.2
NEC, NC, NX, QT, SWC, CC, SC, TW
Macrocixius Matsumura,1914
4
1.6
SWC, CC, SC, TW
Melanoliarus Fennah, 1945
2
0.8
CC, TW
Myndus Stål, 1862
1
0.4
TW
Neocarpia Tsaur & Hsu, 2003
5
2
SWC, CC, TW
Oecleopsis Emeljanov, 1971
14
5.6
NC, SWC, CC, SC, TW
Oliarus Stål, 1862
6
2.4
NEC, NC, SWC, CC, SC, TW
Oliparisca Emeljanov, 2001
1
0.4
QT
Oteana Hoch, 2006
1
0.4
TW
Pentastiridius Kirschbaum, 1868
5
2
NEC, NC, NX, SWC, CC, SC, TW
Reptalus Emeljanov, 1971
6
2.4
NEC, NC, QT, SWC, CC, SC
Semicixius Tsaur & Hsu, 1991
1
0.4
TW
Siniarus Emeljanov, 2007
3
1.2
NC, QT, SWC, CC, SC, TW
Tsauria Kocak & Kemal, 2009
5
2
CC, TW
Mundopa Distant, 1906
1
0.4
TW
Cluster and Ordination
In both the generic and specific taxonomic levels (Fig. 8a, c), the dendograms clearly separate the northernmost regions (Russian Far East, Nei Mongol-Xinjiang and Northeast China regions) from all other regionsand with the similar relationships for Chinese zoogeographical regions: South-Central + SouthWest + North + Taiwan + Qinghai-Tibet. At the species-level the south adjacent China country region appears as sister to all of China. In contrast, at the generic level, this south adjacent China region sister to the central and south Chinese regions. In the northernmost regions, Russian Far East is closer to the Northeast China region at the species level and closer to the Nei Mongol-Xinjiang region at the generic level. In both analyses, the cophenetic correlation coefficient (r>0.8) is high, indicating close agreement between the cluster assignment and the original Jaccard similarity coefficient matrix.
Figure 8.
Dendrograms from UPGMA clustering and NMDS ordination of Jaccard similarity coefficients based on Chinese zoogeographical regions and adjacent areas for Chinese genera (a), (b) and species (c), (d). Abbreviations: NEC, Northeast China; NC, North China; NX, Nei Mongol-Xinjiang; QT, Qinghai-Tibet; SWC, Southwest China; CC, Central China; SC, South China; TW, Taiwan; RFE, Russian Far East; VM, Vietnam, Laos, Thailand, Cambodia, Myanmar, Bhutan, Bangladesh and part of Indian.
The cluster analysis and the NMDS ordination generally showed similar interrelationships among regions (Fig. 8b, d). The stress values of 0.18 (generic level) and 0.30 (species level) demonstrate the accuracy of the projections in the matrix in the 2D ordination space. At the generic level (Fig. 8b), the Nei Mongol-Xinjiang and Russian Far East regions are closely related to each other, and the Northeast China, Nei Mongol-Xinjiang, and Russian Far East regions are clearly separated from the other 7 regions. The Southwest, Central, and South China regions are closely grouped together, and are also related to the North China and Taiwan regions, but the Qinghai-Tibet and VM regions are more separated. At the species level (Fig. 8d), a roughly similar pattern occurs and the Russian Far East is closer to the Northeast China region, but the VM region is clearly separated and more distant from all other regions.
Discussion
Current Chinese diversity and distribution
More than 80% of the species are considered to be endemic to China. The highest endemism is found in Taiwan (69.57%), followed by the Southwest China (39.53%), North China (34.48%) and Central China (33.33%) regions. These figures are consistent with the species richness and endemism patterns observed in other groups, such as aphids (Huang et al. 2008, Gao et al. 2018), leafhoppers (Yuan et al. 2014), or more specifically for planthoppers (Zhao et al. 2020a, Zhao et al. 2020b). For the patterns of distribution, the South China-Taiwan pattern (6.00%), the Central-South China-Taiwan one (4.40%) and the South-Western-South China-Taiwan one (2.40%) are the richer in term of species. This pattern probably results from the past interconnection of the island of Taiwan with the Asian continent during the Quaternary period, when the sea level fell, facilitating the species flow between these areas (Lei et al. 2003, Tang et al. 2006). Its subsequent geographical isolation after the Quaternary period explains its relatively independent pattern of speciation (Gao et al. 2018) and its high endemicity of species.At the tribal level , , and are widely distributed in China, except i the Northeastern China region for the , which is probably a collect artefact as are known to occur in higher latitudes (Bourgoin 2021). With 5.20% of the species, is distributed in the Sino-Japanese - Oriental Region (NC, QT, SWC, CC, SC, TW), and and (6.40% and 2.00% respectively) are mainly concentrated south to the Qingling Mountain-Huai River (SWC, CC, SC, TW). The remaining four tribes [ (0.40%), (0.80%), (1.20%) and (0.40%)] are all distributed in Taiwan.At the generic level, (7.2%) is the most widely distributed genus in China. (2%) is not distributed in the Qinghai-Tibet region. (38%) is not distributed in the Southwest and Northeast China regions, but the genus was reported from the Russian Far East, so it may be a collection bias. In addition, one genus is distributed only in the Tibet region, while 10 genera are distributed only in the Taiwan region. We also found that nearly half of the genera (16 genera, 48.48%) are distributed south of the Sino-Japanese/Oriental boundary.
Biogeographical history shapped Chinese diversity
have a wide range of host plants, including mostly angiosperm Eudicot shrubs: , , , , , etc., but also Monocots as and and tall trees such as Gymnosperm , or Magnolids , etc (Bourgoin 2021). Known from fossil records since the Barremian period, 130MYA (Luo et al. 2020) and probably occurring at least since 200MYA (Urban and Cryan 2012, Song and Liang 2013, Johnson et al. 2018), it is likely that the radiation of angiosperms around 145 MY (Condamine et al. 2020), has greatly influenced the diversification of the major lineages (Labandeira 2014, Szwedo 2016, Luo et al. 2020).More recently, the uplift of the Qinghai-Tibetan Plateau starting in the middle of the Eocene period (45-38 Ma), also had profound effects on the topography and watersheds of East Asia, the aridity of inland Asia, and the Asian monsoon system. These abiotic factors produced a three-stage pattern of species distribution, from high in the west to low in the east (Zhang et al. 2000). The vertical differentiation in plant distribution (Jin et al. 2003), affected their diversity and inceased the richness of local speciation events (Wen et al. 2014, Favre et al. 2015, Ye et al. 2017), and subsequently influenced the species distribution and speciation of the . During the late Oligocene to early Miocene periods (25-15Ma), the expansion of the Tibetan Plateau continued, and the East Asian monsoon and Indian monsoon prevailed in the Asian continent. This resulted in an increase of both temperature and sea levels (Ye et al. 2017), which allowed the northward propagation of fauna and flora. This area was pushed back southwards at the end of the Miocene period (10 Ma) by the uplift of the Hengduan Mountains (Li et al. 2020), which caused the climate to cool (Xie et al. 2019, Yu et al. 2020). Since the middle of the Holocene period (6Ka), rainfall declined and monsoon strength weakened, resulting in a dramatic decrease in precipitation in northern China, which affected the vegetative environment (Zhao et al. 2009, Huang et al. 2012). Quantitative precipitation reconstructions based on pollen collected from northern China indicated that a strong sea-land pressure and temperature gradient caused by strong summer insolation in the northern hemisphere during the early Holocene period (0.14-0.07 Ma) caused enhanced monsoons (Zhao et al. 2009, Cook et al. 2011, Chen et al. 2015, Ge et al. 2017). Obviously the cixiid fauna diversity fluctuated at the same periods along with the diversity of their host-plants. However, without more robust phylogeny studies of , it remains difficult to better infer a more precise biogeographic historical scenario for the family and to link their distribution patterns to any of these important past events.
Biogeographical patterns of Chinese cixiids
Traditionally, the global biogeographical regionalization of China covers both the Oriental and Palearctic realms, which are bounded by the Qingling Mountain-Huai River, around 32–34N in the east of China (Sclater 1858, Wallace 1876, Zhen 1960, Zhang 1999, Cox 2001, Kreft and Jetz 2010, Morrone 2015, Song et al. 2016, He et al. 2017). In 2013, based on its zoological fauna, Holt added a Sino-Japanese realm standing between the Palearctic and Oriental realms, and from west of Tibet to east of the Japanese archipelago. He located the Palaearctic/Sino-Japanese boundary at about 40–41N, and the Sino-Japanese/Oriental boundary at 24–25N in southeast China (Fig. 1). Kreft and Jetz (2013) questioned the validity of this realm because they regarded it as just a biogeographical transition zone between the Palearctic and Oriental realms. According to their taxa etho-ecological characteristics, the Sino-Japanese realm boundaries are generally clustered with the Oriental realm (Kreft and Jetz 2010, Song et al. 2016, He et al. 2017, Gao et al. 2017, He et al. 2018).This result is also observed here for the Chinese divided into two major zoogeographic areas: the Nei Mongol-Xinjiang and Northeast China regions from the rest of China. This boundary corresponds to the Palearctic/Sino-Japanese north boundary and appears to be more well defined than the Palearctic/Oriental boundary. The tribe serves as a landmark for the Palearctic/Sino-Japanese north boundary, while the and tribes are primarily concentrated south to the Qingling Mountain-Huai River point to the traditional Palaearctic/Oriental boundary as proposed by Zhang (2011). and are landmarks for the south Sino-Japonese realm, clustering with the Oriental realm. , , and , which are all distributed in Taiwan, may either indicate the northern limit of older and wider distributions of these tribes or might have resulted from occasional dispersions from neighbouring south regions.At the genus level, the south parts of China cluster with the Indochina region in our analyses, but at the species level all of China forms a unique group. This may be related to the late Eocene uplift of the Himalayas and recent uplift of the Himalayan-Hengduan Mountains in the late Miocene, with a peak before the late Pliocene (Harrison et al. 1992, Sun et al. 2011). These geographical uplifts resulted in the formation of large topographic barriers isolating South China fauna and favorizing recent speciation events and endemism as already shown in several other taxa such as frogs (Che et al. 2010), insects (Ye et al. 2016; Chen et al. 2016), birds (Liu et al. 2016, Cai et al. 2018, Dong et al. 2020), mammals (Ge et al. 2017) and plants (Feng et al. 2013, Favre et al. 2016, Ebersbach et al. 2017, Liu et al. 2017, Ye et al. 2017). Moreover, Quaternary (2.6 Ma) tectonic movements and the influence of the Indian and Pacific monsoons greatly contributed also to the segregation, dispersal and speciation of in southern China and Southeast Asia (Shi et al. 1998, Liang 2003).The South China region is usually included in the Oriental realm in other studies (Zhang 1999, Zhang 2011), but our analysis indicates that for the South China region is closer to the Central, Southwes, and North China region (Sino-Japonese realm). This is consistent with the results of a quantitative analysis of terrestrial mammals in China and adjacent regions by Xiang et al. (2004), where clustering analysis showed the proximity of South China region to Central and Southwest China regions, and they suggested these regions as the South China Division.
Conclusions
This study is the first zoogeographic analysis based on grid cells of in China and adjacent areas, including all the available data for the family. However this dataset has is own limits: 1) the stronger collecting efforts into southern China and taxonomic studies clearly advanced in the Taiwan region because of studies by Tsaur over the past three decades (Tsaur and Lee 1987, Tsaur et al. 1988, Tsaur 1989a, Tsaur 1989b, Tsaur 1990a, Tsaur 1990b, Tsaur et al. 1991a, Tsaur et al. 1991b, Tsaur and Hsu 2003, Tsaur 2009), 2) the limited to very limited knowledge of in countries adjacent to China, despite studies by Distant (1911), Emeljanov (1974), Fennah (1978), Anufriev (1987), Anufriev and Emeljanov (1988), Hoch (2013), Anonymous (2015), 3) it does not take into account host-plants, which are however key factors also affecting the distribution of these obligatory, phytophagous planthoppers, although host-plants and the planthopper species complex are also together affected by other complex topographic and climatic factors embedded in a long dynamic geological process. Accordingly, if the high diversity of Chinese - no less than 8.6% of the current total species richness of the family (Bourgoin, 2021) - is probably related to the high diversity of Chinese biotopes, the figures presented here probably over-estimate the level of endimicity of the fauna in comparison with the adjacent countries.With the current available data, the observed distribution patterns reveals that an intercalary Sino-Japanese realm is recognizable between the Palaearctic and Oriental realms. At the regional level, the South China region clusters more closely with the Southwest, Central and North China regions. Taiwan is clearly separated from the South China region and mainland China, but is more closely related to the Qinghai-Tibet region and Indochina countries. The Central and South China regions are close to each other, but the Qinghai-Tibet region is singularly different. However a much better knowledge of the cixiid fauna in the adjacent countries will be needed in the future for a better evaluation and analysis of the singularity of the Chinese fauna. Additionnaly, a yet to be done phylogenetic analysis of the family will be essential to provide the frame of reference allowing to support any reliable historical biogeography scenario of the evolution, development, and distribution of in China.48 additional species from adjacent areas based on literature and FLOW (Bourgoin, 2021).TablePresence (1) or absence (0) of 48 species in VM (Bangladesh, Bhutan, Cambodia, Laos, Myanmar, Thailand, Vietnam) and RFE (Russian Far East).* BD, Bangladesh; BT, Bhutan; KH, Cambodia; LA, Laos; MM, Myanmar; TH, Thailand; VN, Vietnam.File: oo_611209.docxThe observed material information of checklistspreadsheetExcel version of the observed specimen information for Checklist.File: oo_619349.xlsx
Authors: Kevin P Johnson; Christopher H Dietrich; Frank Friedrich; Rolf G Beutel; Benjamin Wipfler; Ralph S Peters; Julie M Allen; Malte Petersen; Alexander Donath; Kimberly K O Walden; Alexey M Kozlov; Lars Podsiadlowski; Christoph Mayer; Karen Meusemann; Alexandros Vasilikopoulos; Robert M Waterhouse; Stephen L Cameron; Christiane Weirauch; Daniel R Swanson; Diana M Percy; Nate B Hardy; Irene Terry; Shanlin Liu; Xin Zhou; Bernhard Misof; Hugh M Robertson; Kazunori Yoshizawa Journal: Proc Natl Acad Sci U S A Date: 2018-11-26 Impact factor: 11.205
Authors: Ben G Holt; Jean-Philippe Lessard; Michael K Borregaard; Susanne A Fritz; Miguel B Araújo; Dimitar Dimitrov; Pierre-Henri Fabre; Catherine H Graham; Gary R Graves; Knud A Jønsson; David Nogués-Bravo; Zhiheng Wang; Robert J Whittaker; Jon Fjeldså; Carsten Rahbek Journal: Science Date: 2012-12-20 Impact factor: 47.728
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