Pervasive human activities can disrupt invisible facets of biodiversity such as species associations, with potential cascading ecosystem effects (Naeem et al., 1994; Parsons et al., 2022). Human encroachment into natural ecosystems squeezes the spatiotemporal niches of wildlife species (Gilbert et al., 2022), altering the number and magnitude of associations in a community (Burkle et al., 2013), and accelerating species decline and loss (Jones et al., 2018). Humans play a role as ‘super predators’ in shaping the co-occurrence of other species with complicated indirect modifications to multiple interactions between organisms (Gilbert et al., 2022; Moll et al., 2021). Human disturbance, such as chronic landscape modification and acute direct human presence, may significantly alter spatiotemporal distribution of species and fundamentally change the way that species interact (Li et al., 2022a; Sévêque et al., 2020). For example, expanding human footprints have compressed the space and time available for mammals to share by restricting animal movements (Tucker et al., 2018) and increasing wildlife nocturnality (Gaynor et al., 2018). As human and animal activities increasingly overlap in time and space, it is important to assess and quantify the potential for human-induced changes in species association to ecosystem structure and function (Penjor et al., 2022).

Species are not distributed independently of each other; rather, they co-occur in time and space and interact (Tilman, 1994; Wisz et al., 2013). The spatiotemporal co-occurrence of species, termed interspecific associations, provides unique ecological information and has important consequences for ecosystem integrity (Gorczynski et al., 2022; Keil et al., 2021). However, segregated species co-occurrence could be generated by processes such as negative interspecific interaction, distinct environment requirements, and dispersal limitations; similarly, aggregated species pairs may reflect positive interspecific interaction but could also reflect shared environmental preferences (Kathleen Lyons et al., 2016; Song et al., 2020). Although interspecific co-occurrence or avoidance cannot be used to directly estimate species interactions (Blanchet et al., 2020), animals cannot interact if their spatiotemporal niches do not overlap (Gilbert et al., 2022) and strong interactions should be expected to lead to significant associations. Thus, interspecific associations convey key information about interactions between sympatric species (Boron et al., 2023).

Disruptions to the spatiotemporal relationships of species can result in serious ecological consequences including alteration of community structure (Tulloch et al., 2018), upsetting the competitive balance between species (Boron et al., 2023), increasing disease transmission (Hassell et al., 2017), and accelerating local extinction (Fidino et al., 2019). This can significantly distort the distribution of ecological functions that species provide, ultimately influencing ecosystem dynamics (Gardner et al., 2019). Taken as a whole, interactions between sympatric species play a fundamental role in community assembly and are intricately related to ecosystem stability and resilience (Boron et al., 2023). Thus, understanding how interspecific spatiotemporal associations change across human disturbance gradients provides valuable insight into the long-term implications of human impacts on ecosystem function and recovery relevant to biodiversity conservation in the Anthropocene.

Species interactions are known to be context-dependent such that they can vary across space and time, for example along environmental gradients (Chamberlain et al., 2014; Davis et al., 2018; Pellissier et al., 2018; Perrin et al., 2022). For example, gradients in stress are associated with variation in the outcomes of pairwise species interactions (Chamberlain et al., 2014). A key challenge in community ecology is to identify the conditions under which negative and positive species interactions are more likely to occur. For example, the controversial stress-gradient hypothesis predicts that positive interactions should increase as environmental conditions become more severe. Alpine bird communities have been shown to have a higher frequency of positive associations in grasslands with low productivity compared with forests with high productivity (García‐Navas et al., 2021) and Savanah ungulates are more likely to form mixed-species groups in areas where there is predation risk (Beaudrot et al., 2020). Increasingly, major stresses on wild communities derive from human activity, and understanding how species interactions vary in response to human disturbance is pivotal in making robust ecological predictions about biodiversity responses to changing environmental conditions (Dangles et al., 2018; Perrin et al., 2022). However, a review of the literature indicates that the impact of human activity on interspecific interactions of terrestrial mammals remains poorly understood (Boron et al., 2023), particularly in mountain forests where anthropogenic impacts are often pervasive and increasing.

Medium and large-sized terrestrial mammals are key components of mountain forest communities. They play crucial roles in maintaining biodiversity and ecosystem functions (Lacher et al., 2019), and are especially vulnerable to anthropogenic activities (Li et al., 2022b). Here, we set out to investigate the spatial and temporal patterns of occurrence and the interspecific associations within a terrestrial mammalian community along human disturbance gradients. We executed a systematic camera trapping survey spanning 4100 km² of the Yarlung Zangbo Grand Canyon National Nature Reserve in the eastern Himalayas at a total of 322 camera trapping stations, obtaining detections of both wildlife and humans. We classified human disturbances into two groups: human modification (i.e. relating to habitat modifications by humans) or human presence (i.e. referring to the direct presence of humans) disturbances. We employed a context-dependent joint species distribution model (JSDM; Tikhonov et al., 2017) to infer pairwise species associations along human disturbance gradients. We used kernel density distributions of animal diel activity and time between detections of species pairs (time-to-encounter) to compare temporal associations at low and high human disturbances. We consider two hypotheses regarding temporal and spatial interspecific associations. The spatial compress hypothesis posits that human modification of habitats would suppress space available for animals to share (Tilman et al., 2017), resulting in increased positive spatial associations between species. The temporal compress hypothesis postulates that fear of direct human presence would compress diurnal activity in animals (Gaynor et al., 2018), resulting in reduced encounter time (or increased encounter rate) between species (Figure 1). Our study incorporates two different types of human disturbances in the analysis to elucidate the effects of humans on multidimensional (i.e. space and time) species associations. We then consider implications for conservation.

Figure 1

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Our camera traps obtained 10,388 independent detections of 17 focal species (12 carnivores and five ungulates) from 322 stations over 43,163 camera days of effort (Table 1). We documented a number of species and subspecies of conservation concern, including Bengal tiger Panthera trigris, clouded leopard Neofelis nebulosa, and dhole Cuon alpinus. We also captured 2224 independent detections of humans during the survey period.

Species-specific response to habitat covariates

Species varied in their responses to habitat covariates. Nine out of 17 species showed a strong positive response to forest cover (Figure 2, Figure 2—source data 1). Human presence had apparent negative effects on the occurrence of clouded leopard (mean = –0.967, 95% CI=−2.35 to −0.025), taking Budorcas taxicolor (mean = –0.449, 95% CI=−0.935 to −0.071) and red goral Naemorhedus baileyi (mean = –0.806, 95% CI=−0.935 to −0.071), but was apparently positively associated with the occurrence of wild boar Sus scrofa (mean = 0.521, 95% CI=0.045 to 1.093) and golden cat Catopuma temminckii (mean = 0.427, 95% CI=0.165 to 0.804; Figure 2—source data 1). Human modification was apparently negatively associated with the occurrence of dhole (mean = –0.24, 95% CI=−0.493 to -0.027), red fox Vulpes vulpes (mean = –1.588, 95% CI=−2.656 to −0.769), Asiatic black bear Ursus thibetanus (mean = –0.234, 95% CI=−0.42 to −0.056), red goral (mean = –0.665, 95% CI=−1.129 to −0.238) and Mainland serow Capricornis sumatraensis (mean = –0.477, 95% CI=−0.697 to −0.26), but was positively apparently associated with muntjac Muntjac spp. (mean = 3.632, 95% CI=0.946 to 9.999), masked palm civet Paguma larvata (mean = 2.076, 95% CI=1.26 to 3.145), large Indian civet Viverra zibetha (mean = 0.782, 95% CI=0.175 to 1.558) and marbled cat Pardofelis marmorata (mean = 0.689, 95% CI=0.232 to 1.236; Figure 2, Figure 2—source data 1).

Figure 2

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Figure 2—source data 1

Standardized beta coefficients, and 95% credible intervals, for the influence of anthropogenic and environmental covariates on the probability a species used an area during our camera-trap survey in Medog region.

* denotes significant positive and negative effects based on 95% CIs.

https://cdn.elifesciences.org/articles/92457/elife-92457-fig2-data1-v1.xlsx

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Effects of human disturbances on spatial co-occurrence

Out of the 136 estimated pairwise residual correlation coefficients in occupancy, 87 (64 %) were positive at lower human modifications (Figure 3a). At the moderate and higher modifications, the species pairs with positive associations increased to 107 (79 %) and 118 (87%), respectively (Figure 3b and c). At lower modifications, correlation coefficients for 18 species pairs were positive and had a 95 % CI that did not overlap zero (Figure 3—figure supplement 1), and the number increased to 65 in moderate modifications (Figure 3—figure supplement 2) but dropped to 29 at higher modifications (Figure 3—figure supplement 3).

Figure 3 with 3 supplements see all

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Along human presence gradients, 88 pairwise residual correlation coefficients (65%) at lower human presence habitats were positive or close to neutral (Figure 4a). At the moderate and higher human presence habitats, the species pairs with positive associations increased to 115 (85%) and 113 (83%), respectively (Figure 4b and c). The significant positive associations at low, moderate, and higher human presence habitats were 6 (4%, Figure 4—figure supplement 1), 76 (56%, Figure 4—figure supplement 2), and 44 (32%, Figure 4—figure supplement 3), respectively.

Figure 4 with 3 supplements see all

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Effects of human disturbances on temporal co-occurrence

Human presence was associated with significantly increased nocturnality of carnivores (mean = 0.163, 95% CI=0.089 to 0.236, Figure 5a), but showed no significant effects on ungulates (mean = –0.004, 95% CI=−0.031 to 0.023, Figure 5b) and combination of guilds (mean = –0.001, 95% CI=−0.028 to 0.026; Figure 5c). Human modification had no strong effects on nocturnality of any guilds and combination of guilds (i.e. 95% CIs include zero, Figure 5—figure supplement 1).

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Both human modification and human presence were associated with significantly reduced time between detections of pairs (human modification: mean = –1.07, 95% CI=−1.39 to −0.73; human presence: mean = –0.69, 95% CI=−0.92 to −0.46; Figure 6).

Figure 6

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Despite accumulating evidence of widespread impacts of humans on wildlife distribution and activity patterns, our understanding of how different types of anthropogenic pressures reshape species associations remains limited (VanScoyoc et al., 2023). Here, we compared the impacts of human modification (spatial compress effects) and human presence (temporal compress effects) on spatiotemporal associations among threatened terrestrial mammals based on systematic camera-trapping data in an understudied Himalayan landscape. Our results show a strong influence of humans on species co-occurrence patterns. The overall results of the spatiotemporal associations across levels of human modification and human presence suggest that humans are associated with increasing positive spatiotemporal associations among species. Specifically, we detected a higher incidence of positive associations in habitats with moderate and higher levels of human modification and human presence compared to those located in habitats with lower disturbance levels (Figures 3 and 4). On the temporal axis, we detected a significant reduction of pairwise encounter time at increasing levels of human disturbance (Figure 5). Our results, therefore, demonstrate that human disturbance can upset interspecific associations both on spatial and temporal niche dimensions.

Interspecific spatial associations can arise from species interaction, response to environmental covariates, and common dispersal barriers (Blanchet et al., 2020; de Jonge et al., 2021; Poggiato et al., 2021). Our context-dependent JSDM, which accounts for similarities and disparities in species-specific response to habitat covariates considered relevant to medium- and large-bodied mammal species, detected a higher prevalence of interspecific spatial associations in human-disturbed habitats. Humans can have ‘bottom-up’ impacts on animal distribution and associations by land use change and habitat modification (Riggio et al., 2020; Tucker et al., 2018). Human activities can also have top-down impacts on spatiotemporal associations among animals by directly or indirectly changing the landscape of fear (Palmer et al., 2022; Suraci et al., 2019). Previous studies suggest that human disturbance may compress the space and time available for communities to use, resulting in an increased frequency of positive associations (Gilbert et al., 2022; Sévêque et al., 2022). We interpret our results to mean that anthropogenic presence and disturbance reduce available habitats for wildlife, causing a greater predominance of positive associations in anthropogenic landscapes. On the one hand, human presence, such as while grazing livestock and gathering resources, generates landscapes of fear for wildlife (Gaynor et al., 2019; Palmer et al., 2022; Suraci et al., 2019) and may downgrade habitat quality by overexploitation (Filazzola et al., 2020). On the other hand, human modification such as land-use change often constrains the realized niche space of wildlife, restricting animal movement (Smith et al., 2018; Tucker et al., 2018). Overall, our study provides one of the first tests of whether positive spatiotemporal associations between terrestrial mammals increase along gradients of different types of anthropogenic pressures. Our results add to the growing body of evidence that suggests anthropogenic activities reduce available niche space for animals, causing observed positive spatial associations among species (Gilbert et al., 2022; Gorczynski et al., 2022; Murphy et al., 2021).

Identifying thresholds of anthropogenic activity that shift species behavior and co-occurrence will be key to drawing useful inferences from human impact studies and improving our knowledge on when altered associations may lead to reverberating impacts on ecosystems (VanScoyoc et al., 2023; Wilson et al., 2020). We note that the number of species pairs with significant positive associations rapidly increased from lower to moderate levels of human disturbance, but dropped from mean to higher levels of human disturbance (Figure 3—figure supplements 1–3 and Figure 4—figure supplements 1–3). Such patterns are consistent across human modification and human presence levels, indicating a threshold after which the positive effects of human disturbance on species associations were dampened. Human disturbance is an important factor in shaping species distributions (Jones et al., 2018; Samia et al., 2015). Wildlife often has a limited tolerance threshold to human activity (Polaina et al., 2018; Samia et al., 2015; Smith et al., 2019). When human pressures reach levels that preclude a species from occurring at a site (Polaina et al., 2018), prediction of species associations in an anthropogenic context may become uninformative. More work needs to be done to further clarify the mechanisms driving the observed co-occurrence patterns.

Anthropogenic activities can also shift temporal niche of animals (Gaynor et al., 2018; Li et al., 2022a), possibly altering encounter rates among species and trophic dynamics that structure communities (Gilbert et al., 2022; Karanth et al., 2017; Mills and Harris, 2020). Congruent with our hypothesis, we observed a significant reduction of pairwise encounter time at increasing levels of human disturbance, corresponding to more frequent encounters between the pairs (Gilbert et al., 2022). Although we did not find a significant shift in nocturnality at community level, we detected a significant shift to nocturnal activity of carnivores, indicating different sensitivities to human presence among the carnivores and ungulates in our study system. Fear of humans may explain the temporal response of wildlife to human presence. Numerous studies have shown that fear of humans as ‘super predators’ can have suppressive effects on wildlife activity, which may contribute to increased temporal overlap among species (e.g. Sévêque et al., 2022; Suraci et al., 2019).

All in all, our results indicate that both human modification and human presence may rewire species interactions by increasing spatial and temporal co-occurrence. Studies on the impacts of humans on wildlife communities should explicitly account for different types of co-occurring disturbances (Li et al., 2022a). Our camera trapping survey observed that human presence is pervasive even inside this remote protected area in the Tibetan Autonomous Region, indicating that wildlife and their habitats are exposed to frequent human disturbance. Humans exploit resources in protected areas in many ways, including through livestock herding, resource gathering, illegal hunting, and recreation, all of which impact wildlife and their habitats to varying degrees (Harris et al., 2019; Mills and Harris, 2020). Our results demonstrate prevalent disruptions to species co-occurrence patterns from humans. If we are to preserve biodiversity in protected areas, we must work to understand the negative effects on multiple facets of biodiversity from co-occurring anthropogenic pressures. Only then can we design effective mitigation measures.

Although interspecific associations should not be directly interpreted as a signal of biotic interactions between pairs of species (Blanchet et al., 2020; Poggiato et al., 2021), it describes a unique facet of biodiversity, and can provide important insights into the coexistence and persistence of wildlife as well as ecosystem function in settings with anthropogenic activity (Keil et al., 2021; Lai et al., 2020; VanScoyoc et al., 2023). Since interspecific associations often increases with the ecological similarity of species involved (Gorczynski et al., 2022), the tendency of many species towards positive associations with increase in anthropogenic pressures indicates a trend towards homogenization of terrestrial mammal communities. Such human-mediated changes in species co-occurrence patterns can have serious ecological consequences at multiple scales (Gilbert et al., 2022; Gorczynski et al., 2022). For individual species, increased positive associations may affect fitness and population dynamics, increasing local extinction rates (Kuussaari et al., 2009; Parsons et al., 2022). Increase in spatial aggregation among functionally similar species may be indicative of a hidden extinction debt (Kuussaari et al., 2009). At the community level, increased positive associations may depress co-occurrence network complexity and stability, amplify interactions such as predation, and simplify communities with similar traits or co-occurrence patterns (Manlick and Pauli, 2020; Mills and Harris, 2020). Also, increased encounter rates between species may expand disease transmission across communities (Hassell et al., 2017). In addition, wildlife can host a variety of zoonotic diseases (Wicker et al., 2017) and populations unnaturally associated with humans are more likely to transmit pathogens (Jones et al., 2013). We observed that several species such as masked palm civet and wild boar are positively associated with humans in our study area, indicating a substantial overlap between their habitats and human activity. Ongoing habitat modification, livestock grazing, and resource gathering in this region may increase probability of pathogen exchange.

The effects of human disturbance on species associations might be scale-dependent (Gilbert et al., 2022). At present, our sampling design only considers the effects of cumulative human modification and instantaneous local human presence on spatial associations and temporal encounter time at each camera trapping station. We are not able to compare the effects of human disturbance at broader scales (e.g. landscape scale) as such an analysis requires adequate data from hierarchical samplings. Estimating the effects of human disturbance at different spatial scale on species associations is a promising approach. Thus, we encourage future work to further clarify the multiscale ecological effects of human disturbance on species associations.

In conclusion, our study demonstrated that anthropogenic pressures increase spatiotemporal associations of terrestrial mammals from low to mean levels of human disturbances, but the frequency of positive spatial associations dropped from mean to higher levels of human disturbances. Such variations in species associations likely influence the coexistence and persistence of wildlife, with potentially far-ranging ecological consequences. Because terrestrial mammals like carnivores and ungulates play fundamental roles in regulating montane forest ecosystems, the prevalent disruptions to their associations may precipitate biodiversity loss and impair ecosystem function. With increasing human presence and human modification of areas throughout the world, identifying thresholds of anthropogenic activity that shift species relationships, limiting human activity, and increasing landscape connectivity across protected areas may be imperative to maintain interspecific spatiotemporal associations that underpin ecosystem resilience. Moreover, the methods we applied highlight the utility of camera trapping surveys in studying the spatiotemporal relationships among elusive species in settings with anthropogenic activities.

The authors declare that the data supporting the findings of this study are available within the article and its supplementary materials. Occupancy model data (independent detections of wildlife species and human activities histories and model covariates) are openly available in Science Data Bank: https://doi.org/10.57760/sciencedb.11804.

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Author details

1. Xueyou Li

   State Key Laboratory of Genetic Resources and Evolution & Yunnan Key Laboratory of Biodiversity and Ecological Conservation of Gaoligong Mountain, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China

   Contribution

   Conceptualization, Data curation, Software, Formal analysis, Validation, Investigation, Visualization, Methodology, Writing – original draft, Writing – review and editing

   For correspondence

   lixueyou@mail.kiz.ac.cn

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-4705-6082

2. William V Bleisch

   China Exploration and Research Society, 2707-08 SouthMark, Wong Chuk Hang, Hong Kong, China

   Contribution

   Software, Validation, Visualization, Methodology, Writing – original draft, Writing – review and editing

   Competing interests

   No competing interests declared

3. Wenqiang Hu

   State Key Laboratory of Genetic Resources and Evolution & Yunnan Key Laboratory of Biodiversity and Ecological Conservation of Gaoligong Mountain, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China

   Contribution

   Investigation

   Competing interests

   No competing interests declared

4. Quan Li

   State Key Laboratory of Genetic Resources and Evolution & Yunnan Key Laboratory of Biodiversity and Ecological Conservation of Gaoligong Mountain, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China

   Contribution

   Investigation

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0001-7536-5475

5. Hongjiao Wang

   State Key Laboratory of Genetic Resources and Evolution & Yunnan Key Laboratory of Biodiversity and Ecological Conservation of Gaoligong Mountain, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China

   Contribution

   Data curation, Investigation

   Competing interests

   No competing interests declared

6. Zhongzheng Chen

   Anhui Provincial Key Laboratory of the Conservation and Exploitation of Biological Resources, College of Life Sciences, Anhui Normal University, Wuhu, China

   Contribution

   Data curation, Investigation

   Competing interests

   No competing interests declared

7. Ru Bai

   State Key Laboratory of Genetic Resources and Evolution & Yunnan Key Laboratory of Biodiversity and Ecological Conservation of Gaoligong Mountain, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China

   Contribution

   Data curation, Investigation

   Competing interests

   No competing interests declared

8. Xue-Long Jiang

   State Key Laboratory of Genetic Resources and Evolution & Yunnan Key Laboratory of Biodiversity and Ecological Conservation of Gaoligong Mountain, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China

   Contribution

   Conceptualization, Supervision, Funding acquisition, Writing – original draft, Project administration, Writing – review and editing

   For correspondence

   jiangxl@mail.kiz.ac.cn

   Competing interests

   No competing interests declared

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