The Gorkha District is well known as a source of pangolins (Department of National Parks and Wildlife Conservation, 2018). Several individuals have been rescued by the district forest office in this area, which includes community forests, but there is little information on the ecology of pangolins living there. Thus, this study was carried out in Shahid Lakhan Rural Municipality of Gorkha District (27.927032° N, 84.658404° E) in the central region of Nepal (Figure 2). Bhimsen Rural Municipality surrounds this municipality to the north, Gandaki Rural Municipality to the east and Gorkha Municipality to the west. Within Shahid Lakhan Rural Municipality, two areas (Bunkot and Namjung) were selected (Figure 2), covering an area of 44.60 km². The sites studied included community forests, private forests and farmland. The main tree species include Shorea robusta, Castanopsis indica, Schima wallichii, and Acacia catechu. Herbaceous vegetation includes Imperata cylindrica, Hyparrhenia hirta, and Nephrolepis cordifolia. The main castes in the district are Brahmin, Chhetri, Dalit, Gurung, Magar, Newar, Damai, Kami, and Sarki.

Based on a preliminary survey and discussions with local people, areas for the extensive survey were identified. This study was carried out in February 2019. Thirteen transects were established, each 0.5 km long, which were searched for indirect signs (active/old burrows) and any other evidence of pangolin presence (Figure 2). There was a minimum distance of 100 m between adjacent transects in order to avoid overlapping the sites sampled. The transects were carefully chosen to include all habitats based on their relative size. The pangolin burrows were identified based on the characteristics described by Suwal (2011), Department of National Parks and Wildlife Conservation (2018), and Suwal et al. (2020). If a burrow had freshly dug soil, footprints and pangolin feces nearby, it was categorized as an active (new) burrow, whereas burrows with spider webs and dead tree leaves inside were categorized as old burrows (Suwal, 2011). Burrow locations were determined using a handheld Garmin eTrex 30 GPS.

When a (fresh or old) pangolin burrow was found, a circle of 5 m radius with its center at the point where the animal or sign of its presence was located (“presence circle”) was established, following the method of Yahnke (2006) and Bernard et al. (2014), which is the most appropriate in this situation. For each of these circles, an additional similar circle was set up at least 100 m from the former circle in a randomly chosen direction (“comparative circle”), according to Neupane et al. (2022). These plots were samples of average habitat, independent of the presence/absence of the pangolin. Many of the “comparative circles,” therefore, also contained a burrow.

All the “presence circles” and “comparative circles,” which contained burrows indicating pangolin presence, were categorized as “used plots”; those which did not were categorized as “habitat availability plots.” Thus, in total, there were 124 “used plots” and 36 “habitat availability plots.” In all of the “used plots” and “habitat availability plots” seven important ecological parameters were measured (Table 1). These ecological parameters were selected on the basis of previous studies (Wu et al., 2003; Bhandari and Chalise, 2014; Dorji et al., 2020; Acharya et al., 2021; Shrestha et al., 2021). We did not distinguish between old and active burrows in the analyses, as both indicated a recent presence of pangolin.

The households and names of the family heads of all 145 households living within the 3 km buffer zone surrounding the forests in the areas studied were obtained from the village development committee. It was not possible to interview all households because the areas were remote and settlements scattered; therefore, only 87 (60%) of these households were randomly selected for interview. None of them refused to participate in this research.

This survey aimed to determine the people’s knowledge and attitudes to pangolin. It consisted of semi-structured questionnaire (Newton et al., 2008), which was pretested on 5% of the households. The completed questionnaires (see Supplementary material) were followed by face-to-face interviews with the respondents. A random sampling method was used to select potential respondents (Koirala et al., 2012).

Particular households were located with the help of a local field assistant. For the survey, the head of the family was interviewed, and in case of his being unavailable (which does happen and must not exclude the family) the person holding this position in his absence was interviewed. All the respondents completed a questionnaire, which records the demographic (age and gender) and socio-economic status (occupation, education, and ethnicity) of each respondent, which were assigned to different categories, as shown in Table 3.

During the survey, respondents were asked whether they could identify a pangolin based on color photographs, had seen one or any evidence of its presence in the last 2 years, or had knowledge of it and its ecological role. They were also asked about their understanding of pangolin’s legal protection status, their interest in pangolin conservation and the principal threats to its survival. In addition, a survey (n = 9) of the views of members of local government bodies, staff of the divisional forest office, ethnic communities, concerned authorities, police, and traditional healers was completed. During this survey, information on the pangolin’s abundance and major threats to its survival were documented. The questionnaires were recorded in Nepalese or the local language and then translated into English.

In the first analysis, the distribution of pangolin in the area studied was determined by calculating the variance to mean ratio (S²/a), where x is the number of burrows (signs) per transect, a the mean of the x values, and S² = 1n⁢∑(x-a)2 the variance, (Odum, 1971) as follows:

To determine the ecological parameters most frequently associated with the presence of pangolins, a second analysis for each ecological parameter, EP, was done: the percentage frequency of occurrence (P_{EP, Cat}) for each category, Cat, of ecological parameter, EP, was carried out following Pokharel (1996) and Wu et al. (2003):

It must be noted, however, that because the spatial extent of individual categories of the ecological parameters is unknown and it is not possible to measure it and the value of P_{EP, Cat} reflects the spatial extent of each category for each ecological parameter.

In the third analysis, logistic regression was used to estimate the significance of each of the ecological parameters for the selection of individual categories of this ecological parameter by pangolin using all the plots (both “use plots” and “habitat availability plots”) and all data from all the areas studied (Agresti, 2007) in SPSS (IMB Corp, 2015). The dependent variable was the presence/absence of pangolin burrows in plots (set equal to 1 for “use plots” and 0 for “habitat availability plots”), and the independent variables were the values of the seven ecological parameters measured (Manly et al., 2002).

Multicollinearity problems were initially checked for using the variance inflation factor (VIF), which indicated no problems (VIF not >10) (Bowerman and O’Connell, 1990). The P-value of the beta coefficient in the likelihood ratio testing the effect of the independent variable (ecological parameter) at a 5% significance level then revealed whether the corresponding independent variable (ecological parameter) significantly affected pangolin presence or not.

To summarize: the first analysis determined the spatial aggregation of burrows with no consideration of the ecological parameters, the second analysis revealed the relative number of burrows in each category of each environmental parameter but nothing about the preference of pangolin for individual categories and the third analysis, whether there were significant differences in the numbers of burrows recorded for the categories of each environmental parameter.

In the case of social data, respondent variables were noted and used for the analysis of people’s attitude to pangolin conservation. These variables were age (18–35, 36–55, and ≥56 years), gender (male and female) and education (illiterate and literate). The Chi-square test was used to determine whether there are significant differences in the attitudes of the respondents associated with the above variables at the 5% significance level (∝= 0.05).

In total, 124 burrows were recorded: 38 were new and 86 old. The distribution of burrows at different altitudes was clumped, which is indicated by the variance to mean ratio: S²/a = 2.29 > 1. This is also depicted in Figure 2. The highest numbers of burrows were recorded in Rana Khola, Danda Gaun Community Forest, Jugepani Pakha, and Dhurseni and the lowest numbers in Bhalayo Bas Community Forest, Atmare Bhanjyang, Jhagare, and Shikhar Fedi. No burrows were recorded in Magar and Gurung.

Table 1 shows the percentage frequency, P_{EP, Cat}, of pangolin burrows for each of the seven ecological parameters considered. Most of the burrows were recorded at altitudes of between 650 and 800 m a.s.l., in areas with a south-facing aspect, with moderate canopy cover, red soil, gentle terrain, and distance to water <100 m.

The logistic regression model indicates that type of habitat, soil, crown cover, terrain, and distance to water are significantly associated with the distribution of pangolin (Table 2). The highly positive values of the unstandardized beta coefficient, B, in the logistic regression (Table 3) indicate (judging from B in Table 2 and the numbers of burrows in Table 1) that pangolin are strongly associated with: (i) forest and agricultural land; (ii) red and brown soil; (iii) moderate crown cover (26–50%); and (iv) gentle and rolling terrain.

Table 3 lists the sociodemographic characteristics of 87 respondents, in terms of gender, age, occupation, education, and ethnicity.

The questionnaire revealed that of the respondents:

Table 4 lists the responses to the question regarding whether public awareness helps to conserve the habitat of pangolin. There are statistically significant differences in responses based on gender and education. Most males agreed with the statement, whereas female respondents were nearly all neutral. Most literate respondents agreed with the statement, whereas fewer illiterate respondents supported the statement. Responses based on the habitat age were not statistically significant.

Table 5 lists the responses of respondents to the question: do you have an interest in pangolin conservation. There are statistically significant differences based on age and gender. Surprisingly, the numbers of responses of literate and illiterate groups are not statistically significantly different.

Based on the distribution of burrows pangolin distribution is not uniform. Similarly, Suwal (2011), Bhandari and Chalise (2014), and Dhami et al. (2023) report non-uniform distribution of burrows on slopes of all aspects. So, all of the data support the idea that the distribution of pangolin burrows, in the absence of a consideration of the ecological parameters associated with their presence, is clumped.

This study revealed that forest is the preferred habitat of pangolin, with burrows mostly recorded in S. robusta followed by S. wallichii forests. This is in accord with other studies, which report burrows mostly in forested areas (Gurung, 1996; Acharya, 2001; Bhandari and Chalise, 2014; Katuwal et al., 2017: Suwal et al., 2020; Dhami et al., 2023). In forests, termites are very abundant (Ackerman et al., 2009), which could account for the high pangolin occurrence (Swart et al., 1999). In addition, Acharya (2001) report pangolin burrows mostly in S. wallichii and pine forests. So, all of this data support the idea that pangolin prefer forest habitats.

This study revealed that pangolin prefers red soil, which accords with the results of Dhami et al. (2023) reported for Gorkha and Shrestha et al. (2021) for Chitwan, Nepal. In contrast, various studies indicate that pangolin prefers brown soil (Suwal, 2011; Bhandari and Chalise, 2014; Suwal et al., 2020). Heath (1992) for Fujian and Jiangxi provinces in China reports a preference for acidic or yellowish-red soil, which supports the findings of the current study. Thus, it is likely that the “preferred” may be dependent on availability.

This study revealed that pangolins are strongly associated with moderate canopy cover, which is in accordance with the results of Bhandari and Chalise (2014), Dorji et al. (2020), and Dhami et al. (2023). A possible reason for this might be that soils below moderate canopy cover are dry and consist of relatively undecomposed leaf litter and a greater amount of dead twigs and branches (Bhandari and Chalise, 2014), whereas the soil where the crown cover is above 60% is moist with a thick layer of decomposing leaf litter, which is negatively associated with the availability of ants and termites. However, this contradicts Suwal et al. (2020) and Shrestha et al. (2021), who report that pangolins prefer dense crown cover. Therefore, this needs further research.

Wu et al. (2003) report a low number of burrows on slopes greater than 30%, which supports the results presented here of most burrows recorded on moderate slopes. A similar observation is also reported by Bhandari and Chalise (2014) in the Nagarjun Forest in the Shivapuri Nagarjun National Park in Nepal; Sharma et al. (2020c) in Gaurishankar Conservation Area and Ramechhap District in Nepal; and Acharya et al. (2021) in Kavrepalanchok District in Nepal. Pangolin is mostly associated with gentle slopes of between 10° and 30° on which they can easily hunt and feed on ants and termites (Acharya et al., 2021), as well as gentle slopes with an abundance logs that are rich in their preferred prey (Sharma et al., 2020c).

A preference for a certain aspect in different areas might be influenced by climatic conditions, availability of food and degree of human interference (Suwal, 2011). In this study, a high frequency of burrows was recorded on south-facing slopes, which is also reported by Gurung (1996) and Acharya (2001). In contrast, Suwal (2011) reports more burrows on an east-facing slope. A plausible reason for the differing distribution of pangolin burrows on slopes of different aspects in different studies might be that pangolins prefer exposure to direct sunshine, independent of aspect.

Pangolin occurrence is negatively associated with distance to water within a habitat (Katuwal et al., 2017). Thus, similar to the results of the studies by Shrestha et al. (2021) and Dhami et al. (2023), pangolins prefer to live near to a source of water. The reason could be that they need to drink water frequently (Suwal, 2011) in order to regulate their body temperature.

In the current study, burrows were not recorded above 950 m a.s.l., because of the human settlements there and the absence of forest vegetation. A similar observation is also reported by Dhami et al. (2023) for the Gorkha District in Nepal, where the majority of burrows were recorded at altitudes between 450 and 750 m. Bhandari and Chalise (2014) report that pangolins prefer low altitudes, but live mostly at mid-altitudes during winter and Gathorne-Hardy et al. (2001) and Hemachandra et al. (2014) report that termite abundance decreases with increase in altitude, which could be why the number of burrows decreases with increase in altitude. Thus, all the above data supports the idea that pangolin mainly occurs at mid-altitudes.

Very few people knew that pangolins could be a source of income or aware of laws regarding its conservation although it is a protected species in Nepal. According to the fifth amendment of the NPWC Act, there is a provisional fine of NPR 100,000 to NPR 500,000 or jail sentence of 1–10 years or both for the illegal killing, buying and selling and transportation of body parts of protected species (GoN/DNPWC, 2017). In the areas studied, some people used it for meat and medicinal purposes as its meat is considered to be a delicacy and its scales to have curative properties if kept in a cattle shed (Mohapatra et al., 2015). Most of the rural roads that are being constructed are in forests and according to informants they have seen five pangolins in excavator buckets. Thapa et al. (2014) also report that the increase in development works may directly threat the habitat of pangolin. A similar result is that of Katuwal et al. (2017), who report that the construction of footpaths for daily agriculture activities in pangolin habitat directly exposes them to humans resulting in an increase in hunting and poaching.

According to Nash et al. (2016), only 2% of the respondents in Hainan, China, were aware of the medicinal uses of pangolin. However, in Sindhupalchowk District in Nepal, 22% knew that the purpose of the illegal trade in pangolins was their medicinal use (Sharma et al., 2020b). Katuwal et al. (2015) report that 16% of respondents in eastern Nepal believe that pangolin meat can cure gastrointestinal disorders, heart problems, backache, and pain during pregnancy. In addition to its medicinal value, superstitious beliefs are also an important factor in most regions of Nepal because encountering pangolins is believed to bring bad luck (Thapa et al., 2014; Katuwal et al., 2015; Ghimire et al., 2020; Khatiwada et al., 2020).

Anthropogenic activity is the primary threat to pangolin conservation; this finding is similar to that of Baral and Dahal (2022). However, if habitat loss and fragmentation is not greatly reduced or stopped the abundance of pangolins will continue to decrease (Zhang et al., 2022).

Although the majority of respondents in Nepal’s central and eastern regions were aware that pangolin hunting and poaching are illegal (Katuwal et al., 2015; Sharma et al., 2020b), 30.2% had consumed pangolin meat in eastern Nepal (Katuwal et al., 2015). It is mostly unemployed youths aged 16–35 that are involved in the illegal hunting and poaching of pangolins (Katuwal et al., 2015; Ghimire et al., 2020), as it is an easy means of obtaining money (Paudel et al., 2020). Most farmers are unaware of the ecological services provided by pangolin in Nepal (Sharma et al., 2020b). Pangolins are regarded as natural pest control agents, as it is estimated that an adult pangolin consumes more than 70 million insects (mainly ants and termites) annually (Shi and Wang, 1985).

Many local people, on the other hand, claim that burrows reduce crop yield and degrade the aesthetic value of the farmland and report that clusters of burrows can cause landslides during the rainy season (Khatiwada et al., 2020). In Nepal, land conversion for agriculture is expanding (Burton et al., 1989; Paudel et al., 2013), which is accompanied by an increase in the use of insecticides and pesticides (Diwakar et al., 2008; Sharma et al., 2012), potentially reducing the availability of prey for pangolin. Generally there is little or no wildlife conservation in unprotected compared to protected areas (Rodrigues et al., 2004). In the context of Nepal, most pangolin habitat is not in protected areas (Department of National Parks and Wildlife Conservation, 2018), which increases the risk of interaction of this species with local communities.

This study revealed that most respondents did not know anything about pangolins. Suwal et al. (2022) similarly report that the majority surveyed had no knowledge of the ecological, reproductive and behavioral characteristics of this species. Despite the increase in pangolin-related publications in recent years, substantial gaps in knowledge persist in the global literature on various aspects of this species (Heighton and Gaubert, 2021). This includes a dearth of research on pangolin ecology in Nepal, as highlighted by Dhami et al. (2023).

Based on this study, there were significant differences in the extent to which respondents were interested in pangolin conservation depending on their gender and age. Male respondents were more likely to be interested in pangolin conservation, which can be attributed to the fact that males have greater exposure to diverse activities due to better access to education and job opportunities, whereas females are typically confined to household chores and have limited access to such opportunities (Yamamoto et al., 2019). Similarly, old respondents were more interested in pangolin conservation than the young and young adults. Yang et al. (2015) hypothesize that this phenomenon may be attributed to the greater frequency of forest habitat and wildlife encounters in the daily routines of older than younger generations.

This study revealed significant difference across the gender and age groups regarding the question whether public awareness helps to conserve pangolin habitat. Men and educated people may have attended many conservation events hosted by various government and non-governmental organizations, depending on their jobs and obligations. This may help to explain why the male and literate group of participants responded favorably to the aforementioned statement as compared to the female and illiterate group. Thus, pangolin conservation programs should be launched in all the pangolin hotspots, like participatory campaigns in the local community, schools, and radio programs (Goldstein, 2003; Hong et al., 2017) specially designed for females and illiterate people.

Protecting the habitat of pangolins outside protected areas requires people to have a positive perception of endangered animals (Archer et al., 2020). Moreover, conserving these areas as either a national park, community forest or special management unit would safeguard the existence of pangolin in Nepal. Some ecotourism activities can be developed in the area studied, which would help local people economically and improve their attitude towards pangolins (Ngonidzashe Mutanga et al., 2015; Epanda et al., 2019).