Abstract
Thripidae, a member of the order Thysanoptera, has more than 2,200 existing species in the world. Many species of this family are economic pests for agriculture, horticulture, and forestry, whereas the distribution patterns of species diversity at large spatial scales remain poorly understood. We studied the species diversity of Thripidae and the distribution patterns of this family in China, as well as its environmental determinants and geographic determinants, on the basis of 376 species. As a result, 76 endemic species were examined, and Dendrothripinae has the highest percentage of endemic species among 4 subfamilies. The distribution patterns of Thripidae in China are mainly in the Oriental realm; only a few were distributed in the Palaearctic realm. High species richness was concentrated in the South China region, Southwest region, and Central China region. Six of 19 environmental factors were obtained for the relation between species richness and environmental factors. Our study showed that the species distributions of Thripidae in China were mainly influenced by monthly mean diurnal temperature range, Isothermality, SD of temperature seasonal change, Precipitation of the driest month, Precipitation of the wettest month, and Precipitation variation coefficient. The linear relationship between longitude, latitude of the distribution of Thripidae species, and climate factors were explored. Our study analyzed the diversity and distribution of Thripidae in China and provided basic data support for evolutionary biology and conservation of biodiversity.
Keywords: climate, endemic species, environment, model
Introduction
Thripidae is the second largest family in the order Thysanoptera, including more than 2,200 existing species within 289 genera (ThripsWiki 2025). However, it is more abundant than the largest family in temperate regions, whereas this situation is reversed in the tropics. Species of Thripidae are usually tiny, less than 3 mm, but they are widely distributed in all animal fauna in the world. Currently, 4 subfamilies are commonly recognized in Thripidae, Dendrothripinae, Panchaetothripinae, Sericothripinae, and Thripinae. Species of Thripidae are considered to be mainly flower thrips; a great deal of species breed only on leaves, a few are predatory on other small arthropods, and rarely are associated with mosses. Some Thripidae species are key pests with economic effects in agriculture, forestry, and horticulture; they can cause damage by feeding directly but also as a virus vector on plants. Especially some polyphagous species, such as Frankliniella occidentalis and Thrips tabaci, their feeding and virus vectoring have come to involve serious economic losses worldwide (Mound et al. 2022).
Geographic patterns in the distribution of species are crucial to ecology, biogeography, climate change, and the conservation of biodiversity (Gaston 2000, Blanco-Pastor et al. 2019). The spatial distribution pattern of species richness has been examined on a large scale for insects in China (Wei et al. 2016, Zhang et al. 2018, Chen et al. 2020, Jiang et al. 2024). Thripidae species distribution is usually based on species descriptions and records in the world (Denmark et al. 1996, Strassen 2003, Mound et al. 2012, 2017, Cavalleri et al. 2018, Zhang et al. 2018, Okajima and Masumoto 2022).
In China, species composition and fauna distribution characteristics of Thysanoptera in the Qinghai–Tibet Plateau (QTP) revealed that the Sino-Japanese (21.79%) was prominent of Thysanoptera insects in the zoological fauna of the world (Wang et al. 2021). Analysis of relative abundance revealed that fungus-feeding Phlaeothripidae were a common group of soil fauna in the forest litter layer of tropical and subtropical regions (Wang and Zhao 2022). Species diversity and flora characteristics of Thripinae and Sericothripinae revealed that species in the genera Thrips and Neohydatothrips were the most abundant, respectively, with South China as the main distribution area (Hu et al. 2022, 2023), as well as the distribution of Haplothripini (Hu et al. 2019). However, these studies of Thysanoptera were investigated only across small areas or groups, and the environmental factors affecting population dynamics on large spatial scales are limited (Kirk 1997). Therefore, although considerable advances has been made in the study of Thripidae diversity, the relationship between species distribution patterns and the environment remains unknown, especially in China.
Currently, the knowledge of factors affecting the distribution and diversity of Thripidae in China is poorly understood compared to other insect groups. Reliable approaches are required to accurately reveal the diversity of Thripidae across different regions of China. In the present study, we explored the diversity and distribution patterns of Thripidae species in China and assessed the influence of environmental conditions. These findings will improve our understanding of Thripidae species diversity and distributions, and provide a foundation for the conservation of Thripidae species diversity.
Materials and Methods
Species Distribution Data
Species catalog and distribution information of Thripidae from 90% provinces of China were primarily derived from the following sources: collecting records and specimens of Thripidae preserved in the Entomological Museum, Northwest A&F University (NWAFU) (Shaanxi); collecting records from other institutions and academic collections, for example, Institute of Zoology, Chinese Academy of Sciences (Beijing, China), South China Agricultural University (Guangzhou, China), Yunnan Agricultural University (Kunming, China), Australian National Insect Collection (Canberra, Australia); reference records (Han 1997, Wang 2002, Zhang et al. 2012, Zhang 2019, Wang 2020, Feng et al. 2021, Li et al. 2021a, 2021b). Where only administrative locations were given, precise coordinates were obtained using Google Earth according to the location of the administrative center. Recorded data with poorly documented localities (either non-terrestrial or only identified to the provincial/city level) were excluded from analysis. The final database comprised 1,144 records representing 325 Thripidae species and their geographic locations.
Distribution Mapping
China is one of the countries with the richest biodiversity in the world (Xie et al. 2001) and can be generally divided into 7 zoogeographical regions: North-East China Region, Mongolia-Xinjiang Region, North China Region, Qinghai-Tibetan Region, Central China Region, Southwest China Region, and South China Region (Zhang 2011). Data for all administrative units at the province levels (1:4 million) and the Zoogeographical Regionalization of China were from the Chinese Resources and Environment Database (https://www.tianditu.gov.cn/) and the Geographic Data Sharing Infrastructure, College of Urban and Environmental Science, Peking University (http://geodata.pku.edu.cn), respectively. ArcMap 10.8.1 geographic information system software was used to plot the data as XY coordinates on a map of China’s outline. The Chinese territory was divided into 1°×1° grids using the software’s function, and the number of species in each grid was counted to obtain the species richness pattern. We chose this grid size as it is a common standard in Chinese insect diversity studies, matches global environmental dataset resolution (eg WorldClim v2.1), and balances coverage and reliability for our 1,144-record (325 species) dataset.
Environmental Data
In total, the following 19 bioclimatic factors (2.5 arc-minute resolution, 1970 to 2000) were obtained from WorldClim 2.1 (https://worldclim.org/) as the foundational environmental environment dataset: Annual mean temperature (bio1), Monthly mean diurnal temperature range (bio2), Isothermality (bio3), SD of temperature seasonal change (bio4), Max temperature of the warmest month (bio5), Min temperature of the coldest month (bio6), Range of annual temperature (bio7), Mean temperature of the wettest quarter (bio8), Mean temperature of the driest quarter (bio9), Mean temperature of the warmest quarter (bio10), Mean temperature of the coldest quarter (bio11), Annual precipitation (bio12), Precipitation of the wettest month (bio13), Precipitation of the driest month (bio14), Precipitation variation coefficient (bio15), Precipitation of the wettest quarter (bio16), Precipitation of the driest quarter (bio17), Precipitation of the warmest quarter (bio18), and Precipitation of the coldest quarter (bio 19).
Statistical Analysis
The species richness for each grid was determined by counting the number of species within it, and the distribution of Thripidae insects in China was analyzed based on the species richness pattern map. Pearson correlation coefficients were used to test the autocorrelation of the climate factor data by Origin10.1 to avoid the collinearity problem caused by the autocorrelation among the 19 climate factors, and climate data with a high correlation coefficient (|r| > 0.8) were removed in the subsequent analysis.
To further explore the linear relationship between longitude, latitude of the distribution of Thripidae species, and climate factors, each sample station was treated as a distinct point with respect to longitude and latitude; the values from each environmental data were also extracted from each point. The stepwise linear regression analysis was performed using SPSS 26 to examine the associations between climatic factors and the distribution of Thripidae along latitude and longitude. Selected climatic factors (with Pearson correlation coefficients |r| < 0.8, indicating no severe collinearity) were treated as independent variables, while longitude (denoted as Longitude, X) and latitude (denoted as Latitude, Y) served as dependent variables. The “Stepwise” method was adopted, combining forward selection and backward elimination, with inclusion criteria set at P ≤ 0.05 and exclusion criteria at P ≥ 0.10 for the F-test to ensure the model retained only significantly contributing variables.
Results
Species Diversity of Thripidae in China
Among 376 Thripidae species representing 104 genera recorded in China, most genera and all subfamilies of Thripidae were observed, but precise coordinates were available for only 325 species representing 100 genera (around 86% of species and 96% of the genera), and other species were excluded from the distribution analysis because of the absence of precise coordinates. Of the 376 Thripidae species included in this research, 28.99% (109 species) were considered endemic to China (defined as species with no recorded distribution outside China’s geopolitical boundaries, based on global thrips distribution databases and taxonomic records), with obvious variation in the proportion of endemic species across subfamilies (Table 1). Dendrothripinae had the highest proportion of endemic species (11 species, representing 44% of the total endemic species in China) among all subfamilies. Fourteen endemic species in Sericothripinae were sampled, representing 41.12% of all species in this subfamily. Thripinae is the most abundant member, with 76 endemic species reported and take up 27.84% of all species in this subfamily. Eight endemic Panchaetothripinae species were identified, constituting about 18.18% of the total species. One hundred nine endemic species of China were reported totally, but only 3 endemic genera were recorded: Araliacothrips in Panchaetothripinae, Hengduanothrips and Yaobinthrips in Thripinae.
Table 1.
Species diversity of Thripidae in China
| Subfamily | Genera | Species | Endemic species | Percentage of endemic species (%) |
|---|---|---|---|---|
| Dendrothripinae | 7 | 25 | 11 | 44.00 |
| Panchaetothripinae | 17 | 44 | 8 | 18.18 |
| Sericothripinae | 3 | 34 | 14 | 41.12 |
| Thripinae | 77 | 273 | 76 | 27.84 |
| In total | 104 | 376 | 109 | 28.99 |
Distribution Patterns of Species Richness
Thripidae species are found all across China, but as expected in a megadiverse country, their distribution pattern is asymmetric. Species distribution differs significantly between the south (Oriental realm) and the north (Palaearctic realm) of China (Fig. 1), with the majority of species distributed in the oriental realm, and a few in the palaearctic realm. Furthermore, species richness is unevenly distributed among 7 zoogeographical regions (Fig. 2). The high species richness is found in the South China region, followed by the Southwest region and the Central China region, whereas the Northeast region and the Qingzang Region have the lowest species richness. The species richness in the North China region and the Mengxin region is generally moderate, while it shows a relatively high level of species richness in some areas. In terms of the distribution of physical geographical areas, the subtropical and tropical climates are the most concentrated areas of Thripidae species in each climatic zone. The regions with the highest species concentration contained numerous grid cells showing high levels of species richness, mainly focused on south of the Yellow River and east of the QTP, especially including the Southern Qinling Mountains, Southern Yunnan, Yunnan-Guizhou Plateau, Guangdong province, Hainan Island, and Taiwan Island.
Fig. 1.
Distribution of thrips species in China.
Fig. 2.
Geographical division of animals in China.
Environmental and Spatial Associations
Six environment variables were identified through the correlation analysis (Fig. 3), namely, monthly mean diurnal temperature range (bio2), isothermality (bio3), SD of temperature seasonal change (bio4), Precipitation of the wettest month (bio13), Precipitation of the driest month (bio14), and Precipitation variation coefficient (bio15). Five variables were selected in longitude, and the equation fits the interpretation rate 60% at very significant level (P < 0.01) by using stepwise linear regression analysis (Table 2). Precipitation variation coefficient (bio15) was not included in the regression equation, indicating it has a relatively small influence along the longitude distribution. Four variables were selected in latitude, and the equation fits the interpretation rate 92% at a very significant level (P < 0.01). Monthly mean diurnal temperature range (bio2) and Precipitation of the driest month (bio14) were not included in the regression equation, indicating they have a relatively small influence along the latitude distribution.
Fig. 3.
Auto-correlation test of climate factors related to the distribution of Thripidae species in China. bio1, Annual Mean Temperature; bio2, Mean Diurnal Range; bio3, Isothermality; bio4, Temperature Seasonality; bio13, Precipitation of the Wettest Month; bio14, Precipitation of the Driest Month; bio15, Precipitation Seasonality; definitions follow the WorldClim global climate database.
Table 2.
Stepwise linear regression analysis of distribution and climatic factors of Thripidae in China
| Geographical distribution pattern | Stepwise linear regression model | R 2 | df | F | P |
|---|---|---|---|---|---|
| Longitude | Y = −0.659bio3 | 0.250 | 1,221 | 74.16 | <0.001 |
| Y = −0.744bio3 + 0.044bio13 | 0.460 | 1,221 | 94.02 | <0.001 | |
| Y = −0.355bio3 + 0.067bio13 + 0.017bio4 | 0.537 | 3,219 | 85.01 | <0.001 | |
| Y = 0.731bio3 + 0.050bio13 + 0.47bio4 − 3.463bio2 | 0.593 | 4,218 | 79.66 | <0.001 | |
| Y = 0.825bio3 + 0.045bio13 + 0.050bio4 − 3.484bio2 + 0.056bio14 | 0.601 | 5,217 | 65.59 | <0.001 | |
| Latitude | Y = 0.022bio4 | 0.887 | 1,221 | 1735.36 | <0.001 |
| Y = 0.026bio4 + 0.219bio3 | 0.919 | 2,220 | 1252.35 | <0.001 | |
| Y = 0.025bio4 + 0.174bio3 + 0.021bio15 | 0.922 | 3,219 | 861.90 | <0.001 | |
| Y = 0.023bio4 + 0.142bio3 + 0.024bio15 − 0.004bio13 | 0.924 | 4,218 | 662.23 | <0.001 |
Discussion
In the current study, 376 Thripidae species representing 104 genera in China were recorded, but this record possibly underestimates the number of species across this large and environmentally complex area in terms of the thrips' tiny body and hidden living places. Dendrothripinae is differentiated from other Thripidae species by its remarkably elongated metasternal endofurca that usually is related to the saltatorial habit (Mound 1999), and many dendrothripines live on the leaves of tropical forest canopy trees; these factors may have created conditions to evolve separately and form different and endemic species. Larvae and adults of Panchaetothripiane species all feed on mature leaves (Xie et al. 2022); only 8 endemic species out of 44 were reported, and increasing evidence supports a close relationship between the thrips fauna of northern Australia and that of Southeast Asia (Mound and Azidah 2009, Mound and Tree 2009). Wings are crucial for thrips flying and spreading, but some Sericothripiane species exhibit reduction in wing length (Mound and Tree 2009); weak flying ability results in a higher proportion of endemic species in China; 72.6% of Thripidae species were included in Thripinae; of these, 52 species were recognized as economic pest and always have no restricted distribution (Mound et al. 2022). The genetic diversity of these widely distributed species from diverse regions usually has no obvious difference and may be attributed to the strong dispersal ability in the reason of production trade or human activity, coupled with their unique adaptability to the complex environment (Li et al. 2020, Iftikhar et al. 2023, Liu et al. 2023). The rapid, long-distance transport of thrips as stowaways in agricultural produce has greatly expanded the range and enhanced the pest status of numerous species and has facilitated the establishment of short-lived males of bisexual species to survive and reproduce in new regions (Morse and Hoddle 2006).
Thripidae demonstrate greater species diversity compared to any other group of Terebrantia species in China, but many species within this family exhibited a relatively restricted distribution in the current study. Clear differences were found between the northern and southern parts of China with reference to the distribution of Thripidae species, with considerably more species in the south than in the north, as shown by the current compilation of available evidence. The North of China is part of the Palaearctic realm in the zoogeographic regionalization of the world, with the relatively low temperature, but population dynamics data coupled with weather data both reveal a strong positive correlation of the thrips population with high temperature (Rao et al. 2024).
Specifically, Thripidae diversity in China shows core distribution centers concentrated in the South China Region and Southwest China Region, as well as other groups of Thysanoptera (Hu et al. 2019, 2022, 2023). Of these, 109 species were endemic to China, the majority of them from Yunnan, Guizhou, Sichuan, Guangdong, Hainan, and Taiwan. The majority of Thripidae species are herbivorous; the South of China mainly exhibits both tropical and subtropical humid monsoon climates, formed optimal habitats that provide stable thermal and host plant diversity for Thripidae survival. In summary, there are all kinds of types of vegetation cover and physical environments among the different regions and subregions. Inner succession activity, natural conditions, and human disturbance shaped the current Thripidae species distribution pattern.
Previous studies tended to reveal species richness showing a certain pattern along the latitude gradient at large spatial scales (Dunn et al. 2009, Archibald et al. 2010, Stegen et al. 2012). Correspondingly, we observed a marked latitudinal gradient in Thripidae distribution across Chinese subregions, with species richness increasing progressively from north to south, possibly related to the ability of flying of thrips over long distances, due to limited dispersal capacity (Tini et al. 2018). Regression analysis showed that environmental data affected the species distribution along latitude more than longitude, and most of them showed positive effects along longitude and latitude. Temperature emerges as the important predictor among environmental factors. This finding was consistent with previous studies that temperature has a pronounced effect on population dynamics of thrips (Murai 2000, Singh et al. 2017, Karuppaiah et al. 2018). Recent ecological studies demonstrated a positive correlation between thrips population and both the minimum temperature and the morning relative humidity; in general, the hot and dry weather conditions are most favorable for thrips (Rao et al. 2024), and other analysis also reported that the species richness of bacteriophagous tube thrips decreased as temperatures gradually decreased (Wang et al. 2014). The seasonal production in temperate zones of a particularly high diversity of flower-living opportunities facilitated the diversification of flower-breeding Thysanoptera taxa (Zhang et al. 2019). Furthermore, flowers not only provide nutrition but also as habit places; studies of other insect groups also revealed that plant cover has a significant effect on the species diversity, such as ladybugs (Coleoptera: Coccinellidae) (Sushko 2018), stink bugs (Heteroptera: Pentatomidae) (Reisig et al. 2015), and small mammals (Keller and Schradin 2008).
In conclusion, this study presents a comprehensive analysis of Thripidae species diversity and distribution patterns across China by utilizing the extensive occurrence database currently available for these insects in the region. One hundred nine of 376 Thripidae species were found to be endemic in China, and more Thripidae insects were distributed in the south than in the north. The combination of factors is necessary to completely explain the spatial occurrence of Thripidae species, but the current analysis only considered limited relevant factors, and more factors (e.g., percentage of broad-leaved forest) and smaller scales require comprehensive future study. Expanded studies on species occurrence and spatial abundance are needed to further improve and complete the inventory of Thripidae species in China. Meanwhile, effective conservation strategies must be developed to protect the species most susceptible to environmental changes at both regional and global scales.
Acknowledgements
Acknowledgement is given for the data support from the Geographic Data Sharing Infrastructure, College of Urban and Environmental Science.
Contributor Information
Chaorong Meng, Guizhou Provincial Key Laboratory for Tobacco Quality Improvement and Efficiency Enhancement, College of Tobacco, Guizhou University, Guiyang, P.R. China.
Shengjun Jiang, Guizhou Provincial Key Laboratory for Tobacco Quality Improvement and Efficiency Enhancement, College of Tobacco, Guizhou University, Guiyang, P.R. China.
Haixin Qin, Guizhou Provincial Key Laboratory for Tobacco Quality Improvement and Efficiency Enhancement, College of Tobacco, Guizhou University, Guiyang, P.R. China.
Maofa Yang, Guizhou Provincial Key Laboratory for Tobacco Quality Improvement and Efficiency Enhancement, College of Tobacco, Guizhou University, Guiyang, P.R. China; Institute of Entomology, Guizhou Provincial Key Laboratory for Agricultural Pest Management of the Mountainous Region, Guizhou University, Guiyang, Guizhou, P.R. China.
Shimeng Zhang, Guizhou Provincial Key Laboratory for Tobacco Quality Improvement and Efficiency Enhancement, College of Tobacco, Guizhou University, Guiyang, P.R. China.
Author Contributions
Chaorong Meng (Conceptualization [equal], Data curation [equal], Investigation [equal], Software [equal], Visualization [equal], Writing—original draft [equal]), Shengjun Jiang (Investigation [equal]), Haixin Qin (Investigation [equal]), Maofa Yang (Conceptualization [equal], Writing—review & editing [equal]), and Shimeng Zhang (Conceptualization [equal], Data curation [equal], Funding acquisition [equal], Investigation [equal], Writing—review & editing [equal])
Funding
This work was supported by the National Natural Science Foundation of China (No. 32200371), the Guizhou Provincial Basic Research Program (Natural Science) (No. Qian Ke He Ji Chu-ZK [2024] yiban012), and Qian Ke He Ping Tai ZSYS [2025] 028.
Conflicts of Interest
The authors declare no conflict of interest.
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