A total of 2,610 titles were identified from the database searches. An additional 32 articles were identified through other means (including bibliographical search), leading to a total of 2,642 studies. After screening these items to determine whether the inclusion criteria were met, and removing duplicates, a total of 253 articles were assessed for eligibility. The rest of the articles did not have contents for the values but were rather focused on other aspects. Additionally, some of them were duplicates. Of these, an additional 18 were excluded after further screening for whether they contained aspects of socioeconomic benefits. This resulted in a total (n) of 235 studies that were included in our analysis.
Values of bats across regions and countriesA summary of the results from all 235 papers included in the analysis is presented by geographic region (Fig. 1). Each of the different categories of socioeconomic benefits of bats was represented across different geographic areas, although to different extents. It is unclear whether these results reflect differences in reporting and publishing practices, in different regions or different types of activities in each of these categories. The types of activities in each socioeconomic category that were reported from different regions are discussed further below.
Fig. 1Reported values of bats across different continents (n = 235)
Within Africa, records indicate bats were valuable especially in West and East African regions (Fig. 2). They were highly valued for their benefit in form of material and immaterial assets. Notably, sufficient, and nutritious food, income and trade benefits were derived from bats (Fig. 2).
Fig. 2Values of bats across different regions of Africa (n = 75)
In Asia, data indicate bats influenced human livelihoods and well-being more in the South and Southeast areas. They were majorly valued for health aspects of human welfare. The other important value was on the material and immaterial assets which culminated into sufficient and nutritious food (Fig. 3).
Fig. 3Values of bats in Asia (n = 103)
In the Americas, studies indicate values of bats in the South American region were more than that in the North American region (Fig. 4). In both regions, bats were valued for material and immaterial assets. These material and immaterial assets were in the form of fertilizers (e.g., guano), income, sufficient and nutritious food. Meanwhile, in the South American side, benefits centered around health and human communities utilized them to deliver this value through using it as a medicine (Fig. 4).
Fig. 4Values of bats in the Americas (n = 26)
In Europe, records indicate bats were valued majorly in the Southwestern parts (See Supplementary Material 1) and were majorly related with material and immaterial assets (Fig. 5). Through this pathway, it majorly influenced trade and activities to ensure sufficient food. Other benefits included supporting health and were majorly used for medicinal purposes.
Fig. 5Values of bats in Europe (n = 18)
In Oceania, studies indicate bats were majorly associated with material and immaterial components of human well-being. It influenced activities that ensure trade, income generation, transactions and food resources (Fig. 6).
Fig. 6Values of bats in Oceania region (n = 13)
Socioeconomic benefits associated with bats in different areas and across different categories of human welfareMaterial and immaterial benefitsMaterial and immaterial benefits associated with bats were reported more in Africa and Asia compared to other regions (Fig. 7). In these two regions (Africa and Asia), material and immaterial assets were primarily reported in form of supporting sufficient and nutritious food. Other benefits attached to bats within this category and in the two regions included supporting trade, providing fertilizers and income generation (Fig. 7).
Fig. 7Geographical distribution of papers reporting material and immaterial benefits of bats (n = 115)
These values reflect the critical role that bats play in ensuring resource access by human populations. These resources (material and immaterial) are critical in sustaining human livelihoods and well-being across different scales. In Africa, consumption of bats has been recorded in west African countries including Nigeria, Ghana, Benin, Sierra Leone, Cameroon and Côte D’ivoire. In Nigeria, bats form an important source of protein across different communities [22, 28]. Here bats are hunted and consumed. Similar results have been observed in Cameroon [29], Ghana [24, 30,31,32,33], Guinea [22], Côte d’Ivoire [34] and Sierra Leone [35]. Besides hunting of bats for direct benefits including meat, bats were associated with specific ecological services such as enhancing crop production, providing guano as a fertilizer and also suppressing pests. In terms of cash crops, records in Benin and Cameroon indicate bats have significant contribution to enhancing yields of cacao trees through suppressing pests [25, 36]. Bats have also been indicated to be an important source of guano which is used to enhance crop production which is an important ecosystem service [25]. Other material and immaterial benefits associated with bats included generating income through trading activities as observed in Ghana [37].
In Eastern Africa region, bats were reported to be an important source of food, income, guano among others. For instance, anecdotal records from Tanzania and Uganda indicate bat guano to be a valuable resource in ensuring agricultural production. Similar observations were made in Kenya [38]. Besides acting as an important source of guano, bats are hunted and consumed in this region [38, 39]. Meanwhile, in terms of food, bats are an important source of food in form of bush meat within this region [40, 41]. Besides supporting food systems, bats are an important source of income. For instance, in Kenya, bats form an important component of tourism enhancing the welfare of those engaged [38]. Similarly, anecdotal records indicate Python cave (located in Queen Elizabeth National Park in Uganda) to be an important destination for tourists. The tourists often visit this cave to view bats [42]. This generates income not only to those who guide tourists to this site but also contributes to the national revenue. In Tanzania, bats have been noted to form an important part of E-commerce. Notably, bats are turned into souvenirs and sold globally generating income for those engaged in the business and revenue for the country. While we did not find sufficient evidence for the legal status of this business, it appears to be illegal. Consequently, it has potential negative consequences on bat population health as well as public health [43].
In the southern Africa region, the benefit of bats in terms of material and immaterial assets is prominent especially in South Africa and Madagascar. In this region, bats have been noted to act as an important food resource as well as limiting pests. In Madagascar, bats are hunted and consumed as an important source of protein [44,45,46,47]. Meanwhile, in South Africa, bats play fundamental roles in enhancing agricultural production through limiting pest populations. This has been noted in macadamia fields where bats have significantly reduced pests enhancing production [48]. Although scientific reports were not obtained regarding the use of guano in this region, anecdotal records indicate use of bat guano in Mozambique, Zimbabwe, Zambia and Malawi. In Northern Africa, records on the benefits of bats are scanty. However, anecdotal records indicate bats to act as an important source of protein. They are hunted and prepared as a meal.
In Asian countries, bats are majorly valued (in terms of material and immaterial assets) as an important source of food. Other benefits include acting as an important source of revenue, guano, as well as sustaining food production through pollination. In India, bats are hunted and form an important source of protein [49]. Similar observations have been made in Bangladesh [50], Indonesia [51], Malaysia [52], Philippines [53, 54] and Thailand among others [55]. Other benefits include acting as an important source of income and revenue through trade and tourism. Regarding trade, records have been made in Viet Nam where bats get dried and stuck together forming souvenirs [28]. These souvenirs are then sold generating income to those who engage in the business. Meanwhile, in China and Indonesia, bats form an important part of e-commerce [43].
In the Oceania region, material and immaterial benefits have been recorded in the Island nations including Palau, Solomon, and Samoa. In these islands, bats are an important source of food for the local human population. This has been observed in Samoa [56] and Palau islands [30]. The hunted bats are sold generating income. The teeth of bats have also been indicated to be utilized as a form of currency in the Solomon Islands [31]. This is critical in supporting trade enabling access to other needs. In the Oceania region, the bats were majorly associated with sustaining agricultural production through preying on pests. This was indicated in a record by Kolkert and team that indicated bats to contribute significantly to production of cotton [32].
In the Americas, bats have been valued in terms of sustaining agricultural production and generating incomes to those engaged in trade related activities. In terms of agricultural production, Cleveland and colleagues estimated over $700,000 annually as value derived from the pest predation services by Brazilian free-tailed bats [23]. Meanwhile in Brazil, bats have been shown to suppress pests in banana plantations [33]. Similarly, a study by Maslo and colleagues detected DNA of pests in the fecal matter of bats and such pests have a potential to undermine agricultural production [57]. In Chile, bats were shown to limit pest incidences around the vineyards yielding up to $188-$248/ha/year due to bat predation [58]. This reflects the critical role that bats play in sustaining agricultural production. Besides sustaining crop production, bats also generate income for those engaged in trading of souvenirs with bat symbols. This is majorly indicated in anecdotal records. Additionally, in western Brazil, records in the past indicate bats were consumed and formed an important delicacy [36].
In Europe, records indicate bats to be valued largely in terms of sustaining agricultural production as well as generating income and revenue. For instance, in Italy, souvenirs with bat symbols are indicated to be purchased generating income. This has been highlighted on the e-commerce involving bats [43]. Similarly, e-commerce involving bats has been recorded in Germany, UK and Netherlands [43]. In terms of agricultural production, bats have been shown to enhance productivity of livestock in Italy through feeding on pests that would undermine livestock production [59].
Health benefits associated with batsThis benefit mainly linked with medicinal benefits that addressed several ailments and complications. In Asia, South Asia and Southeast Asia were the regions that had more benefits attached to bats in respect to this livelihood component (Fig. 8).
Fig. 8Geographical distribution of papers reporting health benefits of bats (n = 99)
Throughout the world, bats have directly benefited human communities through providing treatment options for different illnesses. These illnesses include asthma, cough, kidney complications, body aches and infertility. Within Africa, the use of bats for managing health-related issues has been observed in West, Central, East and Southern Africa. In West Africa, bats have been used to treat mental illnesses among individuals in Senegal [60]. The head and sometimes whole body of the bat are prepared and consumed to address such ailments [60]. The perception behind this remedy is bats have night-flying ability inferring a symbol of orientation. Therefore, patients with mental illness accordingly have orientation problems and ingestion of certain bat parts can help them recover from the ailment. Although this record was made a long time ago, it may still be practiced up to date and signifies an important socioeconomic value that community members attach to bats. In communities around south western Nigeria, bats are used to treat abdominal pains and infertility [61]. Similarly in Cameroon, bats have been used to address infertility challenges [40]. Similar records have been made in Benin and Ghana among others. Gray literature also indicates bats are used in Nigeria to control baldness.
In the east African region, particularly Kenya, it has been noted to disperse tree seeds providing opportunities for green environment suitable for human health [38]. Meanwhile, in Uganda, anecdotal records indicate community members in the southwestern part of the country eat bats as they perceive them to increase the body immunity [41]. This community was also noted to associate bat consumption with smooth skin among women. Similarly, young children are fed on bat soup to clear off diseases. At times, a tooth of the bat is tied around the waist of the baby and is perceived to clear off diseases. Meanwhile, in the southern African region, health benefits are majorly reported in Mozambique. These health benefits include treatment of cough, baldness and asthma [62, 63].
In Asian region, bats are associated with treatments for asthma, fever, arthritis, liver diseases, cough and tooth ache, among others. In Bangladesh, it is used to treat asthma, fever and arthritis [64]. Bats are also perceived to enhance immunity of people [64]. Meanwhile, in India, bats are noted to provide zootherapeutic benefits including treatment of asthma, cold, cough, tooth ache and liver diseases [65]. They are also utilized to reduce bed wetting [65]. Similar benefits have been observed within communities in Indonesia, Malaysia, Nepal, Pakistan, Philippines, Pakistan and Thailand [45], [99, 100].
In the Americas, Europe and Australasia, health benefits associated with bats appear to have been common in the past but have declined over time. In countries within the Americas including Brazil, bats are pulverized and used in the treatment of asthma in humans [66]. The pulverized product is also used as a contraceptive for livestock [66]. Additionally, it is also used in reducing addiction to alcohol among humans [66]. Within the USA, bat oil was utilized in the past to treat rheumatism [60]. In Latin America, the vampire bats are noted to have saliva with substances that inhibit blood clotting [60]. These substances have been utilized in preventing clotting of blood in patients [60], [102, 103]. This is crucial for patients with acute ischemic stroke [67], [102]. Meanwhile, in European countries, bats played fundamental roles in providing pharmaceutical products [60]. The use of bats for this benefit seems to have faded over time [60]. In Australia, bats are known to be critical in the treatment of asthma [68].
Social, spiritual and cultural benefits associated with batsWithin Africa, this category of benefits associated with bats is more pronounced in West Africa (Fig. 9). This could be explained partly by the differences in social organization across different communities [69]. Social organization within communities influences social capital and other capital assets that are critical in influencing the capability set [54], [107]. Moreover, it varies across space and time with differences in outcomes across the capability set [
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