Discussion
In the present study, we aimed to characterize the spatial genetic structure and demographic history of leopard cats in Taiwan. We identified an interbreeding event that generated a genetic diversity hotspot. We also described a consistent decline in population size that may be associated with increased anthropogenic impacts in Taiwan. The current population structure may indicate which features seem to be barriers to gene flow (e.g. highways, topographical roughness), but natural habitats and main roads at lower elevations may act as movement corridors (Fig. 4). Additionally, we anticipate that climate change will amplify habitat fragmentation in the near future. Moreover, we demonstrated that road-killed samples with a sufficient sample size gathered over an appropriate study area can provide insights into ecological processes and landscape connectivity.
 
Genetic structure and discordance between genetic markers
Our findings supported a correlation between genetic differentiation and landscape features. Whereas our nuclear markers revealed spatial clusters, mtDNA haplotypes exhibited no spatial genetic pattern. Together with the absence of signal of selection on cytb sequences, suggesting cytb sequences alone may not be able to provide biologically meaningful signals for our focal species (Fig. 2d). However, the special genetic analyses using microsatelliates revealed that the predefined populations (i.e., Northern, Central, Southern) demonstrated an obvious and significant genetic differentiation, underlying our hypothesis about the potential barriers of anthropogenic features (e.g., highways) and mountainous ridges to the movements of leopard cats in Taiwan. 
 
Landscape and demographic history shape the current genetic structure of leopard cats in Taiwan
Although our mtDNA data could not provide insights into historical events, by applying a variety of approaches to microsatellites, combining landscape genetic analyses and coalescent methods (Epps & Keyghobadi, 2015), we revealed associations between current landscape features and population structures with historical evolutionary events also contributing to the genetic profile of leopard cats.
 
Our analysis of colonization history revealed a divergence between the Northern and Southern populations 800 years ago, followed by a secondary contact 200 years ago. We infer that this secondary contact generated the genetic diversity hotspot and the pattern of admixture observed in the Central population from the previously isolated populations. For example, human activities and the development of natural habitats provide opportunities for migration and result in secondary contacts (Grabenstein & Taylor, 2018). Similarly, the intended development and population growth of humans may force the historically differentiated Northern and Southern populations to contact again in the region of the current Central population within the past 200 years.
We found evidence of bottleneck events in the past 200 years in all three populations, coinciding with rapid human development in Western Taiwan (Barclay, 2015; Wu, 2004). Although the exact cause of this change in demographic history remains unclear, historical climate fluctuations and human activities are potential contributing factors. Dramatic demographic changes of mammals in other regions have been attributed to climate variations and human disturbances (Hsiung & Sunstein, 2006). Habitat loss, deforestation, and changes in land cover for residential or commercial use may deteriorate the persistence of several species in Taiwan, including leopard cats (Chang et al., 2012; Chen et al., 2016; Hwang et al., 2010). Consequently, the drastic population decline event followed by human-mediated migration may be attributed to the distribution of leopard cats nowadays that strictly limited to few regions in Western Taiwan.
 
Gene flow in a heterogeneous matrix with human disturbance
Despite inconsistencies between results generated using PCA-derived or Nei's genetic distance and ResistanceGA and RCM approach, elevation and roads appeared among the most critical landscape features to shape landscape genetic connectivity in Taiwanese leopard cats. Contrary to our expectations, resistance to movement increased with elevation, even though natural forests dominate mountainous regions. This outcome may be explained by a preference for forest edges, which provide easy access to refuge in forests and prey in agricultural landscapes for leopard cats and European wildcats (Hartmann et al., 2013; Rajaratnam et al., 2007). Tracking studies conducted in Taiwan also suggested that leopard cats prefer densely vegetated natural habitats and avoid areas with human activities (van der Meer et al., 2023). An alternative explanation can be derived from the concept of the energy landscape, whereby landscape complexity influences animals to adopt the most energetically efficient movement patterns (Wilson, Quintana, & Hobson, 2012). We observed a positive correlation between resistance and roughness (a proxy of topological complexity), but resistance only increased significantly at high roughness levels and higher elevations (>1,000 m). Thus, landscape complexity alone does not fully explain why elevation hampers connectivity at lower elevations. We propose a third potential explanation based on the typical physiological intolerances to large temperature variations of tropical organisms (Janzen, 1967). We found that the distribution of leopard cats in Taiwan was negatively correlated with minimum temperature (Fig. S9), indicating that low temperatures may limit their expansion and movement. Unlike their relatives in temperate areas, tropical and subtropical leopard cats are typically restricted to lowlands (Rajaratnam et al., 2007; Ross et al., 2015). Moreover, elevation as a significant barrier has been reported for diverse taxa in tropical regions (Monteiro et al., 2019). It is noteworthy that using RCM and DPCA genetic distance, the best-supported model included Land use and Human density, suggesting the putative influences of anthrophonic factors may also be a profound factor that shapes the movement of leopard cats, as revealed in the studies of other felids (Hartmann et al., 2013). 
 
Linear features such as paved roads and highways are artificial structures known to have a significant impact on gene flow in many species (Clark et al., 2010; Kuehn et al., 2007; Robinson et al., 2012), and highways have been identified as physical barriers that impede movement and lead to a decline in genetic diversity (Epps & Keyghobadi, 2015). Huge artificial linear features may also act as behavioural barriers (Kimmig et al., 2020; Laundré, Hernández, & Ripple, 2010). Similarly, we inferred that the avoidance of highways of leopard cats in Taiwan may be driven by a perceived risk associated with high-speed traffic and strong illumination. In contrast, we found that unfenced roads presented low resistance, serving as corridors that facilitate movement. Such roads are often located in areas with lower human activity and traffic volume than highways. In mountainous regions, roads are typically situated in valleys at lower elevations and are surrounded by natural forests, making them suitable for leopard cat movement (Chen et al., 2016; Pei, 2008). Although vehicle collisions pose a significant mortality risk for leopard cats in Taiwan (at least 50 road-kills recorded between 2012 and 2017) (Chen et al., 2019), they may still use roads as corridors to avoid other human activity areas. Similar patterns have been observed for foxes, which prefer sites of low human density and activity rather than completely preventing human constructions (Adkins & Stott, 1998). These findings highlight how different linear landscape features influence connectivity and can contribute to our understanding of dispersal processes in vagile carnivores.
 
Conservation implications
Our genetic study complements previous ecological studies revealing that direct and indirect impacts of anthropogenic features have compromised the sustainability of Taiwanese leopard cat populations. Although there is currently no evidence of severe genetic drift that may significantly affect the genetic diversity among leopard cat populations in Taiwan, human-mediated landscape features have substantially contributed to genetic differentiation among its populations. Despite no prospect of significant loss of diversity in the near future, our species distribution models predict that climate change may further endanger the viability and connectivity of Taiwanese populations, restricting suitable habitats to northern and mountainous regions with rugged topography, which the high elevation and complex topography are supposed to act as profound resistance to movement and gene flow (Fig. 5c; Fig. S9). The mitochondrial marker revealed extremely low genetic diversity in the leopard cat populations of Taiwan among related populations in Far East Asia (Ito & Inoue-Murayama, 2019; Ko, An, & Eo, 2022; Teng et al., 2022). Moreover, the distinct Taiwanese haplotypes indicate limited gene flow between peripheral island habitats and continental core populations (Patel et al., 2017; Tamada et al., 2008). The evolutionary potential to adapt to changing environments depends on the balance between genetic variation and drift (Brown, 1984; Holderegger, Kamm, & Gugerli, 2006). However, the low genetic variation resulting from founder effects and lack of new genetic components increases the risk of local extinctions in island populations facing extreme climate change (Johnson, Adler, & Cherry, 2000; Wood et al., 2017).
The critical conservation management actions for Taiwanese leopard cats should focus on maintaining neutral and adaptive genetic diversity, population fitness, and evolutionary potential in the face of habitat fragmentation and impending climate alterations. Efforts should be made to reduce adverse human activities that drive genetic differentiation in the island's leopard cats, such as establishing corridors to facilitate gene flow between fragmented habitats. Protected areas can be designated as areas of high dispersal potential to minimize disturbance from human activities and predation by free-roaming domestic dogs. To mitigate fragmentation and reduce the risk of road collisions, artificial underground or aboveground corridors can be implemented along highways in areas with a high population density of leopard cats (Beier & Noss, 1998). Lastly, to counteract the negative effects of drift and inbreeding caused by isolation and climate change, establishing corridors connecting southern and northern areas should be a high priority to prevent the loss of genetic diversity and reduce the risk of local extinctions. Our study provides valuable insights into how landscape features shape the evolutionary processes and future distribution of a locally endangered felid on a subtropical island, offering fundamental ecological information for its conservation.
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Statements & Declarations
Funding
This work was supported by the National Science and Technology council under project IDs 106-2313-B-002-048, 107-2313-B-002-004, and 108-2313-B-002-001.
 
Competing Interests
The authors have no relevant financial or non-financial interests to disclose.
 
Author contribution
Y.T.J. designed and supervised the project. J.C.P., Y.H.L., D.J.L., M.T.C., and P.K.J.C. collected the samples and provided valuable feedback. C.H. and L.W. performed laboratory experiments and analyzed the mitochondrial data. P.W.S. performed statistical analyses of demographic history, landscape genetics, and niche modeling using the input from C.H. and L.W. P.W.S. and C.H. drafted the manuscript. P.C.L. assisted with data interpretation, provided valuable feedback, and revised the manuscript. All authors have read and approved the final version of the manuscript.
 
Data Accessibility
DNA sequences are available at Supporting Information and NCBI GenBank (PbTaiC1: accession number OR126345; PbTaiC2: accession number OR126346; PbTaiC3: accession number OR126347). All microsatellites and cytb data genotyped in this study were deposited on Figshare (DOI: 10.6084/m9.figshare.24188403)
Supporting Information