Carnivores play key roles in maintaining ecosystem structure and function, as well as ecological processes (Ripple et al., 2014). Apex carnivores are often classically specialized hunters occupying top trophic positions that work to suppress the number of herbivores and mesocarnivores through predation, competition, and trophic cascades (Ripple et al., 2014; Newsome et al., 2017; Ritchie and Johnson, 2009). Understanding how sympatric carnivore species coexist in natural ecosystems is a central focus in community ecology and biodiversity conservation (Chu et al., 2017). Human activities and climate change are causing large-scale habitat loss and fragmentation, posing significant challenges to carnivore interspecies relationships (Li et al., 2021; Manlick and Pauli, 2020; Ripple et al., 2014). Such relationships are becoming increasingly intricate, leading to greater research efforts aimed at determining the strategies that facilitate coexistence despite intense competition (Kuijper et al., 2016; Smith et al., 2017).

The competitive exclusion principle dictates that species with similar ecological requirements are unable to successfully coexist (Hardin, 1960; Gause, 1934). Thus, carnivores within a guild occupy different ecological niches based on a combination of three niche dimensions, i.e., spatial, temporal, and trophic (Schoener, 1974). Spatially, carnivore species within the same geographical area exhibit distinct distributions that minimize overlap in resource use and competition. For example, carnivores can partition habitats based on habitat feature preferences and availability of prey (de Satgé et al., 2017; Garrote and Pérez de Ayala, 2019; Goldyn et al., 2003; Strampelli et al., 2023). Temporally, differences in seasonal or daily activity patterns among sympatric carnivores can reduce competitive interactions and facilitate coexistence. For example, carnivores can exhibit temporal segregation in their foraging behaviors, such as diurnal versus nocturnal activity, to avoid direct competition (Finnegan et al., 2021; Nasanbat et al., 2021; Searle et al., 2021). Trophically, carnivore species can diversify their diets to exploit different prey species or sizes, thereby reducing competition for food resources. For example, carnivores can exhibit dietary specialization to optimize their foraging efficiency and minimize competitive pressures (Steinmetz et al., 2021). Currently, research on the niche partitioning of sympatric carnivores primarily focuses on one or two dimensions, with little attention paid to all three (Alexander et al., 2016; Karanth et al., 2017; Li et al., 2022a; Santos et al., 2019; Shao et al., 2021; Strampelli et al., 2023; Tsunoda et al., 2020; Vilella et al., 2020). This limitation restricts the multi-scale understanding of coexistence mechanisms among carnivores.

In recent years, camera trapping and DNA metabarcoding technology has been widely used in wildlife monitoring and research. Camera trapping enables the monitoring of elusive species in remote areas (Boitani, 2016; Palencia et al., 2022), and has become a helpful tool for accumulating large amounts of time-recorded data that can provide detailed information surrounding animal behavior and activity patterns, which is widely used to investigate species-interspecies relationships (Frey et al., 2017). For example, Li et al., 2019, used camera trap data to conclude that temporal segregation is a key mechanism for promoting the coexistence of tigers (Panthera tigris) and leopards (Panthera pardus). DNA metabarcoding provides a noninvasive molecular tool that is more accurate than traditional dietary analysis methods, such as microhistology (Deiner et al., 2017; Newmaster et al., 2013). DNA metabarcoding technology has been applied to many studies and has offered profound insight into the ecology, conservation, and biological monitoring of rare and endangered species (Deagle et al., 2019; Kartzinel et al., 2015). For example, Shao et al., 2021, found that dietary niche partitioning promoted the coexistence of species in the mountains of southwestern China based on DNA metabarcoding. The combination of camera trapping and DNA metabarcoding can result in robust data for exploring the mechanisms of coexistence surrounding carnivore guilds.

The Qilian Mountains constitute a biodiversity hotspot with one of the richest carnivore assemblages in the world (Di Minin et al., 2016). To understand the competition and coexistence among different carnivore species in this area, we explored the habitat use, activity patterns, and prey item composition of sympatric carnivore species comprised of apex carnivores and mesocarnivores across Qilian Mountain National Park using camera trap data and DNA metabarcoding data. Based on theories surrounding resource partitioning and niche differentiation, as well as studies on interactions and coexistence among carnivorous species (Haswell et al., 2018; Linnell and Strand, 2000), we hypothesized that differentiation along one or more niche axes is beneficial for the coexistence of the carnivorous guild in the Qilian Mountains. We expected that spatial niche differentiation promotes the coexistence of large carnivores in the Qilian Mountain region, as they are more likely than small carnivores to spatially avoid interspecific competition (Davis et al., 2018). Mesocarnivores may coexist either spatially or temporally due to increased interspecific competition for similar prey (Di Bitetti et al., 2010; Donadio and Buskirk, 2006). Dietary niche differentiation may be a significant factor for promoting coexistence between large and mesocarnivore species due to differences in body size (Gómez-Ortiz et al., 2015; Lanszki et al., 2019).

Our study addresses, for the first time, the coexistence patterns of carnivore species present in Qilian Mountain National Park across multiple dimensions of niche partitioning. This work substantially contributes to current understanding of carnivore guilds and offers helpful information for biodiversity conservation at a regional scale. Moreover, our study provides important insights into the potential mechanisms of niche partitioning among sympatric carnivores, particularly intra- and interspecific relations between apex carnivores and mesocarnivores. Specifically, we found that the overall trend of spatial overlap across carnivores is relatively low, that apex carnivores overlap in time and diet, that mesocarnivores showed a high degree of dietary overlap, and that there was substantial similarity in diel activity patterns between apex carnivores and mesocarnivores. These results indicate that carnivores with similar ecological traits foster co-occurrence by adjusting their daily activity patterns and using differing food resources to minimize competitive interactions.

We found dietary and temporal overlap among apex carnivores, showing that spatial partitioning is responsible for their successful coexistence in this area, consistent with our hypothesis. Wolf and snow leopard had the highest dietary overlap and prey similarity between apex carnivore pairs in this study, showing that the avoidance of space and time plays an important role in their coexistence. Recent evidence proves that habitat preference facilitates the coexistence of wolf and snow leopard (Shrotriya et al., 2022). Their hunting strategies may be impacted by their habitat selection. Solitary snow leopards are more suitable for hiding in habitats with features that favor ambush predators, while wolves hunt in packs (Shrotriya et al., 2022). It is clear that wild ungulates (e.g. blue sheep) constituted the primary proportion of wolf and snow leopard diet, followed by small mammals such as plateau pika, Himalayan marmot (Marmota himalayana), and woolly hare (Lepus oiostolus). In addition, livestock consumption also contributed to the high degree of overlap in their diets (Wang et al., 2014). This supports the optimal foraging theory, in which large predators preferentially select food resources that provide maximum benefit (Brown et al., 1999), but also showed that greater competition for resources is likely to occur between wolf and snow leopard due to their use of the same prey species in this area. This may be especially true in times of habitat stress when resources are poor.

Snow leopard and Eurasian lynx had the highest temporal overlap between apex carnivore pairs in our case, showing that spatial and dietary partitioning facilitate their coexistence. Eurasian lynx is considered an opportunistic predator, and its prey varies among different regions with its primary dietary resource being ungulates and small mammals. For example, Eurasian lynx showed a strong preference for brown hare Lepus europaeus in Turkey, edible dormice Glis glis in Slovenia and Croatia, and chamois Rupicapra rupicapra or roe deer Capreolus capreolus in Switzerland (Mengüllüoğlu et al., 2018; Krofel et al., 2011; Molinari-Jobin et al., 2007). Varied prey selection may be related to sex, age, population density, and season (Mengüllüoğlu et al., 2018; Odden et al., 2006). Our results show that woolly hare make up the majority of the Eurasian lynx diet, followed by blue sheep. Woolly hare is mainly distributed in shrubland, meadow, desert, and wetland, while blue sheep tend to choose highly sheltered areas, close to bare rocks and cliffs as habitat. Therefore, prey preferences among snow leopard and Eurasian lynx also contribute to spatial avoidance.

Mesocarnivores had substantial overlap in diet, underscoring food resources as a primary competitive factor that necessitates spatial and temporal partitioning for successful coexistence. This finding aligns with previous research indicating that mesocarnivores use temporal and spatial segregation to reduce competition and the probability of antagonistic interspecific encounters (Ferreiro-Arias et al., 2021; Li et al., 2022a). Differences in habitat preference may lead to spatial niche partitioning among mesocarnivores (Wang et al., 2022). In addition, species can adjust temporal periods of behavior to respond to environmental change, competition, and predation risk (Gallo et al., 2022; Finnegan et al., 2021; van der Vinne et al., 2019). Pallas’s cat hunts during crepuscular and diurnal periods and inhabits meadow habitat with greater prey abundance (Anile et al., 2021; Greco et al., 2022; Ross et al., 2019). In contrast, red fox is primarily nocturnal and occupies diverse habitats depending on prey abundance (Goldyn et al., 2003; Pandolfi et al., 1997; Reshamwala et al., 2022). Tibetan fox is a diurnal hunter of the Tibetan plateau, preferring shrub meadow, meadow steppe, and alpine meadow steppe (Gong and Hu, 2003). It is worth noting the substantial overlap in diet between Pallas’s cat and Tibetan fox. The dietary overlap between the two was extremely high, with a Pianka’s value close to 1. Dietary analyses showed that pika contributed to more than 85% of their collective diets, with 90% of Pallas’s cat diet being pika. Pika may be an optimal prey item in the area because of size and year-round activity (Ross, 2009). Previous studies have shown that the Pallas’s cat and Tibetan fox are specialist predators of pikas (Harris et al., 2014; Ross, 2009). However, specialization on pika is facultative in that Pallas’s cat and Tibetan fox can select other prey items when pika availability is low (Harris et al., 2014; Ross, 2009). This was observed in our study, even though dietary diversity was low.

Apex carnivores and mesocarnivores exhibited considerable overlap in time overall, showing that spatial and dietary partitioning may play a large role in facilitating their coexistence. As confirmed by previous research, kit foxes (Vulpes macrotis) successfully coexisted with dominant carnivores by a combination of spatial avoidance and use of alternative resources (Lonsinger et al., 2017). Differences in body mass may play a crucial role in minimizing dietary overlap, effectively reducing interspecific competition between apex and mesocarnivores. Of exception in our study, however, was wolf and red fox, who exhibited more dietary overlap, showing that temporal and spatial avoidance may promote their coexistence. As canid generalist-opportunist species, the wolf and red fox consumed similar prey, albeit the red fox may have obtained livestock and ungulate species via scavenging or by preying on very young individuals (Hacker et al., 2022). In addition, the wolf and red fox had different peak activity times, suggesting temporal segregation as a potentially strong driver of coexistence. Our occupancy model also revealed that mesocarnivores prefer habitats that coincide with human disturbance, possibly contributing to spatial niche differentiation with other apex carnivore species like the wolf. Our findings echo other recent research which supports the idea that red foxes can coexist with wolves by exploiting a broader ecological niche (Shrotriya et al., 2022).

Our study highlights the effectiveness of combining camera trapping with DNA metabarcoding for detecting and identifying both cryptic and rare species within a sympatric carnivore guild. This integrated approach allowed us to capture a more comprehensive view of species presence and interactions compared to traditional visual surveys, whereas it is important to acknowledge the challenges associated with this technique, including the high costs of equipment and the need for specialized training and computational resources to manage and analyze the large volumes of sequence data. Despite these challenges, the benefits of this combined method in improving biodiversity assessments and understanding species coexistence outweigh the drawbacks. However, several restrictions remain for this research. The first limitation involves differences in samples sizes. Although the scat samples of Tibetan fox were relatively low, the accuracy of DNA metabarcoding in informing species presence in diet ensures that data are informative and thus still important for species conservation management decisions (Hacker et al., 2022). Second, the methodology of foraging (e.g. predation or scavenging) and the condition of the prey item (e.g. age or size) cannot be identified in dietary studies (Hacker et al., 2022). Pika is a prime component of diet among mesocarnivores, especially in the diet of the Pallas’s cat and the Tibetan fox. We surmise that the simultaneous dependence on pika led to partial overlap in spatial and activity patterns, resulting in increased potential competitive interactions. Due to the lack of spatial and temporal analysis of Pallas’s cat in our study, further monitoring is needed to develop a comprehensive conservation plan. Despite these limitations, our study provides a foundation from which future studies interested in niche partitioning among carnivores along spatial, temporal, and dietary dimensions can be modeled.

In summary, our study has shown that the coexistence of carnivore species in the landscapes of Qilian Mountain National Park can be facilitated along three niche axes, with spatial segregation being the most pronounced of the three. Apex carnivore species tended to overlap temporally or trophically, mesocarnivore species had high dietary overlap with each other, and apex carnivore and mesocarnivore species displayed similarity in temporal use. Pika, blue sheep, and livestock were found to make up a large proportion of carnivore diet. Resource competition between wolf and snow leopard and the interspecific competition between Pallas’s cat and Tibetan fox were strong in this area. Based on the findings presented above, we recommend targeted efforts for enhanced protection and management in several key areas. First, greater efforts are needed to protect habitat, including the establishment of habitat corridors and optimizing grassland fence layouts to safeguard migration routes. Second, resource competition should be carefully monitored between snow leopards and wolves, as well as between Pallas’s cat and Tibetan foxes. More attention is needed for pika at our study site considering the role they play in the conservation of Pallas’s cat and Tibetan fox populations. Previous poisoning campaigns targeting pikas were initiated out of concern for grassland degradation (Smith and Foggin, 1999). Recent research underscores the importance of pika population health and habitat for the distribution of Pallas’s cats (Greenspan and Giordano, 2021). It’s crucial to note that poisoning campaigns targeting small mammals may be incredibly dangerous for mesocarnivores feeding on them due to secondary poisoning, necessitating caution in such conservation strategies. Further, vigilance regarding the preponderance of smaller prey in predator diets is vital, as this may indicate severe loss of larger prey, which will increase the risk of interference competition (Steinmetz et al., 2021). Third, efforts should focus on restoring vulnerable wild prey populations, strengthening grazing area management, and supporting livelihoods of herders to mitigate livestock predation and consequent human-wildlife conflicts. Regular patrols of protected areas and core habitats are essential, along with educational outreach to conservation staff and herders. Our study corroborates and complements the findings of prior studies on these species and their coexistence mechanisms, offering insights crucial for wildlife conservation in the region.

All data generated or analysed during this study are included in the manuscript and supporting files.

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Author details

1. Wei Cong

   Ecology and Nature Conservation Institute, Chinese Academy of Forestry, Key Laboratory of Biodiversity Conservation, State Forestry and Grassland Administration, Beijing, China

   Contribution

   Investigation, Writing – original draft, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0009-0005-6071-9564

2. Jia Li

   Institute of Ecological Conservation and Restoration, Chinese Academy of Forestry, Beijing, China

   Contribution

   Funding acquisition, Investigation, Writing – review and editing

   Competing interests

   No competing interests declared

3. Charlotte Hacker

   Department of Biological Sciences, Duquesne University, Pittsburgh, United States

   Contribution

   Investigation, Methodology, Writing – review and editing

   Competing interests

   No competing interests declared

4. Ye Li

   Ecology and Nature Conservation Institute, Chinese Academy of Forestry, Key Laboratory of Biodiversity Conservation, State Forestry and Grassland Administration, Beijing, China

   Contribution

   Investigation

   Competing interests

   No competing interests declared

5. Yu Zhang

   Ecology and Nature Conservation Institute, Chinese Academy of Forestry, Key Laboratory of Biodiversity Conservation, State Forestry and Grassland Administration, Beijing, China

   Contribution

   Funding acquisition, Investigation

   Competing interests

   No competing interests declared

6. Lixiao Jin

   Ecology and Nature Conservation Institute, Chinese Academy of Forestry, Key Laboratory of Biodiversity Conservation, State Forestry and Grassland Administration, Beijing, China

   Contribution

   Investigation

   Competing interests

   No competing interests declared

7. Yi Zhang

   Ecology and Nature Conservation Institute, Chinese Academy of Forestry, Key Laboratory of Biodiversity Conservation, State Forestry and Grassland Administration, Beijing, China

   Contribution

   Investigation

   Competing interests

   No competing interests declared

8. Diqiang Li

   Ecology and Nature Conservation Institute, Chinese Academy of Forestry, Key Laboratory of Biodiversity Conservation, State Forestry and Grassland Administration, Beijing, China

   Contribution

   Funding acquisition, Methodology, Project administration

   Competing interests

   No competing interests declared

9. Yadong Xue

   Ecology and Nature Conservation Institute, Chinese Academy of Forestry, Key Laboratory of Biodiversity Conservation, State Forestry and Grassland Administration, Beijing, China

   Contribution

   Data curation, Investigation, Project administration, Writing – review and editing

   For correspondence

   xueyadong334@163.com

   Competing interests

   No competing interests declared

10. Yuguang Zhang

    Ecology and Nature Conservation Institute, Chinese Academy of Forestry, Key Laboratory of Biodiversity Conservation, State Forestry and Grassland Administration, Beijing, China

    Contribution

    Data curation, Funding acquisition, Investigation, Methodology, Project administration, Writing – review and editing

    For correspondence

    yugzhang@sina.com.cn

    Competing interests

    No competing interests declared

    "This ORCID iD identifies the author of this article:" 0000-0001-9801-8556

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