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Distribution of tick species in mainland China
We compiled a database comprising 7344 unique records on geographic distributions of 124 known tick species, including 113 hard tick species in seven genera and 11 soft tick species in two genera, together with 103 tick-associated agents detected in either ticks or humans, which were recorded in 1134 counties (39% of all counties in the mainland of China) (Supplementary Fig. 1 and Supplementary Note 1). The most widely distributed tick genus was Dermacentor (in 574 counties), followed by Heamaphysalis (570), Ixodes (432), Rhipicephalus (431), Hyalomma (298), Argas (90), Ornithodoros (38), Amblyomma (37), and Anomalohimalaya (5) (Supplementary Data 1 and Supplementary Figs. 2‒10). At the species level, D. nuttalli, Ha. longicornis, D. silvarum, Hy. scupense, and R. sanguineus were each found in >200 counties, followed by R. microplus, I. persulcatus, I. sinensis, I. granulatus, and Hy. asiaticum that were each detected in 100‒200 counties (Supplementary Data 1). We identified 19 predominant ticks that were detected in more than 40 counties, including five Ixodes species, four Heamaphysalis, four Dermacentor, three Rhipicephalus, two Hyalomma, and one Argas tick species. Forest and meadowlands are the major vegetation types for these 19 tick species, accounting for a median of 46.4% (IQR: 40.0%‒68.9%) of their habitats (Supplementary Data 1).
The abundance of tick species varies substantially across the seven biogeographic zones which are defined by climatic and ecological characteristics (Fig. 1)18,19. Tick species are most abundant in Central China, South China, and Inner Mongolia–Xinjiang districts, hosting 61, 57, and 50 tick species, respectively (Supplementary Data 2). Eight prefectures reported ≥20 tick species, three in Xinjiang Autonomous Region of northwestern China, two in Yunnan Province of southwestern China, and one in each of Gansu, Hubei, and Fujian provinces of northwestern, central, and southeastern China, respectively (Fig. 1). Most genera except for Amblyomma were found in northwestern China, particularly in Xinjiang Autonomous Region. In contrast, less tick diversity was observed in northeastern China, which only harbors Ixodes, Heamaphysalis, and Dermacentor (Supplementary Figs. 2‒10).
Fig. 1: Tick species richness (circles) at the prefecture level in seven biogeographic zones in mainland China from 1950 to 2018.

I = Northeast district (NE), II = North China district (N), III = Inner Mongolia–Xinjiang district (IMX), IV = Qinghai–Tibet district (QT), V = Southwest district (SW), VI = Central China district (C), and VII = South China district (S). Source data are provided as a Source Data file.

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Risk mapping and risk factors for 19 predominant tick species
The ecological modeling results for the 19 predominant tick species showed highly accurate predictions, with the average testing area-under-curve (AUC) ranging from 0.83 to 0.97 (Table 1) and the testing partial AUC ratio ranging from 1.30 to 1.78 (Supplementary Tables 1‒5), indicating decent predictive power. The ecoclimatic and environmental variables that were predictive of the geographic distribution of the ticks differed among the species, even for those in the same genus (Fig. 2f, Supplementary Tables 1‒5). Temperature seasonality and mean temperature in the driest quarter were the two most important drivers, contributing ≥5% to the ensemble of models for 14- and 12- tick species, respectively, followed by elevation contributing ≥5% to the models for seven tick species (Fig. 2f, Supplementary Tables 1‒5). The same predictor, however, may drive the risk in different directions for different tick species (Supplementary Figs. 11‒29). For example, a high temperature in the driest quarter was associated with a high probability of presence for I. granulatus and R. haemaphysaloide but with a low probability for I. persulcatus and Ha. longicornis (Supplementary Figs. 11, 13, 16, 22).
Table 1 The average testing areas under the curve (AUC) of the BRT models at the county level and model-predicted numbers, areas and population sizes of affected counties for the 19 most prevalent tick species in China.
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Fig. 2: Clustering of tick species based on their ecological features and spatial distributions at the county level.

Panels a‒e indicate the spatial distribution of the five clusters (clusters I‒V). The boundaries of the seven biogeographic zones are shown as black solid lines. The dendrogram in panel f displays the clusters I‒V of tick species. The features used for clustering are three quantities associated with each predictor in the BRT models. Two of the three quantities were displayed in panel f to indicate the possible level of ecological suitability: relative contributions (colors in ascending order from yellow to red) and the standardized median value of the predictor (numbers in the heatmap) among counties with tick occurrence (numbers 1‒4 indicate the position of this median in reference to the quartiles of this predictor among all counties). Source data are provided as a Source Data file.

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The model-predicted high-risk areas of the 19 tick species were much more extensive than have been observed, 31‒520% greater in the number of affected counties, 14‒476% larger in the size of affected geographic area, and 25‒556% larger in the affected population size (Table 1, Supplementary Figs. 30‒34). Ha. longicornis was predicted to have the widest distribution that potentially affected 588 million people in 1140 counties, followed by I. sinensis and R. microplus that affected 363 and 350 million people in 630 and 678 counties, respectively (Table 1). High-risk areas of these three tick species collectively covered nearly all densely populated areas in China, mainly provinces in the central, eastern, southern, and southwestern China (Supplementary Figs. 30(b), 31(a), and 32(b)). R. sanguineus, and R. haemaphysaloides each affected more than 200 million people. D. nuttalli, I. crenulatus, Hy. asiaticum, Ar. persicus, and D. daghestanicus ticks were the top five tick species affecting the largest areas at the scale of 2.0‒3.8 million km2 (Table 1).
Ecological clustering of tick species
Based on the ecological similarity represented by the environmental and ecoclimatic predictors, the 19 tick species were grouped into five clusters with clear patterns of spatial aggregation (Fig. 2). D. nuttalli and D. silvarum constituted cluster I that covered the vast region in northern (including northeastern and northwestern) China. This cluster stretches over biogeographic zones I‒IV characterized by middle to high elevations, shrub grassland, strong seasonality in temperature, relatively low temperature in the wettest quarter (often also the warmest quarter), and low precipitation in the driest month (Fig. 2 and Supplementary Figs. 23, 24). Ha. longicornis, Hy. scupense, and R. sanguineus were grouped into Cluster II which was mainly found in biogeographic zones II, III, and VI, featuring the landscape of shrub grassland and irrigated or rainfed croplands at low-middle elevations (7% (Table 2). SFTSV ecologically prefers regions at low to moderate elevations ( 7%). High risks of TBEV were flagged by low to medium elevations ( More