Biodiversity Conservation and Use

The Concept, Connotation and Significance of Cultural Keystone Species in Agricultural Heritage Systems

  • MIN Qingwen , 1, 2, * ,
  • YANG Xiao 1, 2 ,
  • DING Lubin 1, 2
  • 1. Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing 100101, China
  • 2. University of Chinese Academy of Sciences, Beijing 100049, China
* MIN Qingwen, E-mail:

Received date: 2021-07-31

  Accepted date: 2021-10-15

  Online published: 2022-01-08

Supported by

The Strategic Priority Research Program of the Chinese Academy of Sciences(XDA23100203)


As a new type of heritage, Agricultural Heritage Systems (AHS), represented by Globally Important Agricultural Heritage Systems (GIAHS) designated by Food and Agriculture Organization of the United Nations (FAO) and Nationally Important Agricultural Heritage Systems (NIAHS) designated by some countries’ Ministry of Agriculture, are typical Social-Ecological Systems (SES), which usually are rich in biodiversity, traditional knowledge, resource utilization technology and outstanding cultural landscapes. Cultural Keystone Species (CKS) are defined as the culturally salient species that shape the cultural identity of a people in a major way. CKS can be used as a prominent tool for the synergistic conservation of SES biology and culture, and to promote the overall enhancement of system functions. This paper summarizes a review of the definition of the CKS and its application in SES conservation. According to the characteristics and protection needs of AHS, this paper defined the CKS in AHS as: “Composites of biological resources and cultural practices, which have a significant impact on the stability of local society and culture systems, contribute to the achievement of AHS’ conservation goals.” Based on this definition, we analyzed the significance of the identification of CKS in AHS. First of all, CKS help to quickly identify the key elements of AHS. Secondly, CKS can promote community participation in the conservation and development of AHS. In addition, the identification of CKS has a significant role in food and livelihood security, biodiversity conservation, traditional knowledge and technology transmission, social organization maintenance, and cultural landscape maintenance in AHS, which helps to achieve the conservation goals of GIAHS and/or NIAHS.

Cite this article

MIN Qingwen , YANG Xiao , DING Lubin . The Concept, Connotation and Significance of Cultural Keystone Species in Agricultural Heritage Systems[J]. Journal of Resources and Ecology, 2022 , 13(1) : 51 -60 . DOI: 10.5814/j.issn.1674-764x.2022.01.006

1 Introduction

In the field of Social-Ecological Systems (SES), the coordinated protection of biodiversity and cultural diversity has gained international consensus. In 1988, the Belem Declaration clearly pointed out that there is an “inseparable link” between biodiversity and cultural diversity (UNESCO, 1988), in which the concept of “biocultural diversity” was put forward to solve the crisis of diversity loss in SES for the first time. After this perspective was put forward, it was quickly introduced into ethnobotany research by ethnobota-nists (Pei, 2011a). It was proposed that “taking the co-evolution of plant diversity and cultural diversity as the main line, to study the cultural behavior and practice process of human utilization of plants and its influence on plant species and ecosystem” (Long et al., 2009). Since then, in order to study the interaction between human and plants more deeply, ethnobotanists have discovered that just as “ecological keystone species” play a role in ecosystems, some species occupy an important position in the cultural systems of local communities, such as religious stories, folk festivals, songs, languages, and food. Such species have been called “Cultural Keystone Species (CKS)” (Garibaldi and Turner, 2004; Cristancho and Vining, 2004), and the CKS are defined as organisms that have played an important role in the stability of local cultural systems. These culturally salient species that shape the cultural identity of a people in a major way, are often reflected in the fundamental roles these species have in diet, materials, medicine, and/or spiritual practices (Garibaldi and Turner, 2004).
Since then, a number of researchers have explored in depth of the effects of CKS on local SES structure for a particular CKS. For example, Singh et al. (2015) studied Paisang (Quercus griffithii) as a CKS sustaining local livestock, conserving more than 20 local livestock species and crop varieties, and providing important livelihood values to the local communities. Jacques-Coper et al. (2019) point out that Andean Condors in the Andean region are national symbols of governance, represent national unity, contribute to the evolution of social relations, and are also embedded in the spiritual beliefs of the inhabitants of the local communities, and closely related to their worldview. Chen and Akamine (2021) found that Garcinia subellipticas in Japan are mostly planted around villages and have an important windbreak function, forming a unique rural landscape. Butler et al. (2012), in a study on traditional ecological knowledge (TEK) and fisheries management in the Torres Strait, Australia, considered Turtles (Chelonia mydas) and Dugong (Dugong dugon) as local CKS. But because of overfishing, the populations of these two species have declined dramatically and the government has requested their protection. Because of their unique cultural values, local residents have a high level of acceptance of this initiative, facilitating the distribution of power and community co-management of resources in conservation actions. Another study on kahawai (Arripis trutta) similarly pointed out that in New Zealand fisheries management, local residents not only conveyed their beliefs to young people through kahawais fishing, but also established an ecological view of not over-harvesting from nature, a concept that also promoted local community support for kahawai as a conservation program (Maxwell et al., 2018). From these examples, it can be seen that “CKS” play important role in research areas such as ecological restoration (Ann, 2009), community co-management (Freitas et al., 2020), and TEK conservation (McMillen, 2008).
As a new type of heritage, Agricultural Heritage Systems (AHS), represented by Globally Important Agricultural Heritage Systems (GIAHS) designated by Food and Agriculture Organization of the United Nations (FAO) and Nationally Important Agricultural Heritage Systems (NIAHS) designated by some countries’ Ministry of Agriculture, are typical SES. GIAHS Initiative aims at discovering and conserving typical and threatened traditional agricultural systems around the world. According to FAO (2002), GIAHS are defined as “Remarkable land use systems and landscapes which are rich in globally significant biological diversity evolving from the co-adaptation of a community with its environment and its needs and aspiration for sustainable development”. Since the day it was proposed, the AHS has been regarded as a typical SES (Parviz, 2009; Li, 2015), facing the problem of coordinated protection of biodiversity and traditional culture. However, most of the previous studies on AHS failed to take into account the relationship between biodiversity and traditional culture (Zhang et al., 2016; Jiao et al., 2021). The protection of AHS through the identification of CKS not only promotes the synergistic conservation of biodiversity and traditional culture, but also helps in the identification of key elements and the conservation of priority objects in heritage sites. However, there are few studies on CKS in China, and its application in AHS is relatively rare. Based on a review of the definition of the CKS, and its application to SES conservation, CKS and its connotations are defined here in the context of AHS. Finally, the significance of CKS identification for the conservation of AHS is explained. In conclusion, this paper aims to provide a perspective and methodology for further research on the synergistic conservation of biology and culture, and to provide a basis for the conservation and development of AHS in the future.

2 Concept and connotation of CKS in AHS

2.1 Concept of CKS in AHS

It is generally believed that the concept of CKS was inspired by “ecological keystone species” (Garibaldi and Turner, 2004), although there is a key difference in emphasis between them. “Ecological keystone species” refers to those species that play key roles in the structure and function of the ecosystems and are vital to the integrity of the ecosystems (Paine, 1969). The “Cultural Keystone Species” focuses more on the impact on local cultural systems and refers to plants and animals that have an irreplaceable role in the formation and development of local community culture. From the current research, Garibaldi and Turner’s definition of CKS is more frequently cited, but there are other researchers who have different opinions.
After this concept was put forward, Ellen noticed that there were many similarities between “CKS” and the “ethnobiological keystone species” concept that he put forward earlier (Ellen, 2006). The ethnobiological keystone species was defined as “species are crucial to the maintenance of the entire anthropogenic environments, subsistence systems and ways of life”. In this definition, Ellen assumes that the ethnobiological keystone species is equivalent to a single species. Platten and Henfrey (2009) raised objections. As human ecologists, they believe that the concepts and connotations of CKS should be defined from a systematic perspective. Rather than a biological species, the CKS should be regarded as a composite of natural organisms and social culture with interrelated functions. However, Freitas et al. (2020) believed that we should use the more comprehensive term “culturally important species” (CIS), considering that some species may play an overriding role in people’s culture yet are not necessarily irreplaceable or indispensable to the culture's survival. They also point out that the extinction or degradation of CKS can affect the transmission of traditional knowledge and the continuation of traditional practices. Thus, there is no clear definition of CKS. The only thing that is clear from either perspective, is that in the overriding principle of the definition of CKS, it has a significant impact on the local sociocultural system. Therefore, the definition of CKS in AHS should also follow this principle, and should be consistent with the main characteristics and conservation objectives of AHS.
As a new type of heritage, rooted in a long cultural tradition and long-term practical experience, AHS are a typical socio-ecological system in which human activities interact with the natural environment and involve the whole body (Li, 2015). The AHS contain many important and complex subsystems, such as economy, biology, technology, culture and landscape. These subsystems are interrelated and influence each other. For example, the abundant local biological resources not only provide people with various types of food or raw materials to support their livelihoods and economic systems, but also is the material carrier of local traditional culture. In turn, the traditional knowledge and culture accumulated by local people over a long period influence the way they use biological resources and shape the outstanding cultural landscapes. It can be seen that AHS encompasses the long-standing habits in the use of agricultural organisms by local communities and their interactions with local culture (Min, 2021), which are the core content of CKS. In addition, according to the five criteria for identifying GIAHS given by FAO (2002), AHS must have the following characteristics: food and livelihood security, agro-biodiversity, local and traditional knowledge systems, cultures value systems and social organizations and landscapes features. Therefore, the conservation of AHS should also focus on these five standards, including: maintaining the function of agricultural production as an important source of livelihood for local farmers, maintaining traditional crop types and land use patterns, inheriting traditional agricultural production knowledge and technology systems and maintaining the stability of local social organizations and agricultural landscapes.
It can be seen that AHS have outstanding biodiversity and traditional cultural diversity. Especially in the long- standing human-land interaction, local people have accumulated a great deal of traditional knowledge, such as the identification of edible wild plant resources and the knowledge of traditional medicine. Based on the characteristics and conservation requirements of AHS, we believe that CKS must be closely related to local traditional cultural knowledge and be able to play an important role in promoting the preservation of the core elements and overall structure of AHS. Therefore, its loss can have a significant impact on one or more aspects of AHS, such as food and livelihood security, conservation of biodiversity and ecological functions, transmission of traditional knowledge systems, perpetuation of cultural values and social organization, and maintenance of local landscapes. In addition, we incorporate Platten and Henfrey’s view that CKS should not be directly equated with a particular biological species when defining CKS in AHS (Platten and Henfrey, 2009). Because although CKS are based on biological resources, their contribution to the system structure also depends on a series of cultural factors, such as traditional festivals, religious beliefs, language vocabulary, social norms and values. In summary, CKS of AHS refers to composites of biological resources and cultural practices, which are closely linked to local traditional cultural knowledge, have a significant impact on the stability of local society and culture systems, and contribute to the achievement of the conservation goals of GIAHS and/or NIAHS.

2.2 Connotation of CKS in AHS

According to the above definition of CKS in AHS, the connotation of CKS contains both natural and cultural attributes. The natural properties part refers to the organisms contained in the CKS. In the process of identifying the CKS, this part can be divided according to the method of biological classification, called biological classification units, which determines the rank of the identified organisms. What we want to point out is that the organisms that can be considered as CKS should be divided into wild and cultivated categories in AHS. When identifying wild plants or animals, the basic unit of biological classification units is the species. For example, the animals associated with cultural totems in the Aruqorqin Grassland Nomadic System include gray wolf (Canis lupus L), plum deer (Cervus nippon), and celestial eagle (Accipiter gentilis), as well as the fishy grass (Houttuynia cordata Thunb) and spiny coriander (Eryngium foetidum) in Honghe Hani Rice Terraces System. However, the basic unit of biological classification units needs to be further subdivided when identifying cultivated plants or animals. This is because with cultivated organisms it is more difficult to reveal the interactions between organisms and cultures only at the species level. It has been demonstrated that different subspecies or different varieties of cultivated organisms play very different roles in the local cultural systems (Yang et al., 2019). For example, in the Honghe Hani Rice Terraces System, the same rice species is used, but glutinous rice and red rice have very different effects on the cultural system (He et al., 2020; Ding, 2021). Similar phenomena are found in other AHS, such as in the Diebu Zhagana Agriculture-forestry-animal Husbandry Com-posite System, where fern pigs play a more important role in the cultural system than other domestic pigs (Yang, 2019).
The cultural attributes of CKS refer to the cultural practices such as traditional knowledge which are related to biology. There are many different types of cultural practices in AHS, which will be expressed in the form of production, life, beliefs, customs, etc. In the process of identification, in order to clearly understand the value and status of CKS, we believe that we can draw on the concept of cultural practice units proposed by ethnobotanists Winter and McClatchey (2009) in their study of the changing relationship between plants and people, which refers to different types of cultural activities. According to the relationship between cultural practices and local peoples’ lives, cultural practices can be divided into production cultural practice units, living cultural practice units and spiritual cultural practice units. Production cultural practice units can be divided into agricultural calendar, agricultural production technology, and handicraft culture. Life cultural practice units can be divided into food culture, medical culture, clothing culture, architectural culture, and transportation cultures. Spiritual cultural practice units can be divided into festivals customs, religious beliefs and values (Table 1).
Table 1 Division of cultural practice units in AHS
First classification Second classification Examples of representative cultures of AHS
Production cultural
practice units
· Agricultural production technology Rice-fish-duck three-dimensional breeding technology; Water resources management technology, such as water diversion woodcut and water diversion stone carving
· Agricultural calendar Twenty-four Solar Terms; Kaiyangmen, Saguzhong
· Handicraft culture Sculpture craft; Ceramic craft; Ware making
Life cultural practice units · Food culture Long Street Banquet; New Eating Festival; Duck Catch Festival; Frozen Fish Festival
· Medicine culture Application of traditional Chinese medicinal materials
·Dress culture Tie dyeing; Silk reeling; Textiles; Embroidery
· Architectural culture Mushroom House; Dry Bar House structure
· Traffic culture Use of animal drawn vehicles
Spiritual cultural practice units · Festival customs Sacrifice to Heaven, Stars and Fire; Fish Lantern Dance; Shiyuejie
· Religious beliefs Multi-god worship; Ancestor worship
· Tradition values The values of harmony between man and nature and respect for nature

3 The significance of CKS for AHS

3.1 Easily identifying the key elements of AHS

The key elements of AHS are the composites of material and non-material elements representing the typical characteristics of the heritage system, maintaining the long-term stability of the system structure, and promoting the multiple value realizations of the system (Li, 2021). The identification and analysis of key elements is important to understanding the structure and function of AHS. However, because of the complexity of the components, the identification of key elements has become one of the priority areas requiring attention in the current conservation of AHS (Min, 2020; Li, 2021).
According to the current research, there are few relevant studies that address the key elements of AHS. In terms of identification methods, Li (2021) used the combination weight method and the crowd proportion method to identify the key elements of the Honghe Hani Rice Terraces Systems and Shexian Stoned Dry Terraces System. However, the methods and indicators he used are still relatively rough, and the results are not very accurate. For example, the key elements of Honghe Hani Rice Terraces System are identified as food crops, terrace building technology, Hani traditional diet and so on. These results are not only ineffective, but also fail to consider the interactions between various elements.
If we focus on protecting a limited number of CKS, the system is easier to manage than trying to identify and protect all of the key elements in the system. It is also more appropriate to the reality of the limited human, material, and financial resources invested in current development. Therefore, if we adopt the perspective of CKS for the identification and management of key elements, we can not only effectively identify different biological classification units and cultural practice units, but also help to sort out the interaction between local agrobiodiversity and local traditional culture. This approach can achieve dynamic conservation and sustainable development of the AHS with half the effort.

3.2 CKS is a unique perspective for studying biocultural diversity

Since the 1980s, biologists, ecologists and environmentalists have realized the impacts of society, culture, and economy on biodiversity (Pilgrim et al., 2009). The role of culture in protecting biodiversity has gradually become one of the core issues of biodiversity research. Some scholars pointed out that the real driving force of the major threats to biodiversity is the power of cultural values that people choose to develop economically, rather than the economy itself (Pei, 2011b). Afterwards, more and more studies have pointed out that biodiversity and cultural diversity are facing common threats, and they are co-evolving and influencing each other (Davidson et al., 2012). More scholars have begun to believe that biodiversity helps maintain the health of natural ecosystems and cultural diversity helps maintain the health of social systems. In addition, the joint protection of the biodiversity and cultural diversity can improve the overall resilience of SES. Therefore, the conservation of biodiversity and cultural diversity should be integrated, from which the concept of biocultural diversity is derived (Pilgrim et al., 2009). In 2010, the Convention on Biological Diversity (CBD) and United Nations Educational, Scientific and Cultural Organization (UNESCO) jointly launched the “Joint Programme on the Links between Biological and Cultural Diversity”, indicating that the concept of biocultural diversity had become an international consensus (Agnoletti and Rotherham, 2015).
At present, the research content of biocultural diversity mainly focuses on traditional knowledge, cultural landscapes and the interaction between biology and culture (Mao et al., 2017), which integrates the dual perspectives of the natural and human sciences. However, there are still many methodological barriers to the study of biocultural diversity because of the long-standing opposition between the research paradigms of the natural and human sciences (Pilgrim et al., 2009). Although it also implies the interaction of humans and nature, the focus of CKS is on finding a limited number of organisms and associated cultures that are crucial to the maintenance of cultural systems (Fig. 1). As a result, ethnobotanists have paid a great deal of attention to the concept of CKS as a prominent perspective and approach to the study of biocultural diversity (Cristancho and Vining, 2004; Garibaldi and Turner, 2004; Gaoue et al., 2017).
Fig. 1 Relationships between CKS and biocultural diversity and related concepts

3.3 Promote the enthusiasm of local people to participate in co-management

AHS involve many stakeholders, such as the government, local communities, tourists, and researchers. As users and inheritors, local peoples are one of the most important subjects of the heritage systems. First, the local peoples, as the main cultural inheritors, have a deeper understanding of the traditional culture of the heritages. Secondly, they provide labor, services and other products for the conservation and development of the heritages. What’s more, local peoples, as users of rich local biological resources, have long practiced a harmonious man-earth relationship, and they are indispensable for the preservation and transmission of the heritages.
CKS, which are species or varieties closely related to the traditional culture and knowledge in AHS, usually play important roles in the productive life of the local people. At the same time, in the process of identifying CKS, the frequency of use by local peoples, the number of occurrences in the traditional culture and the status in the local language are important measurement indicators.
As a result, local peoples are more willing to accept CKS and have a better understanding of the role that the CKS play in maintaining the stability of their local socio-cultural systems. If we begin our conservation efforts by focusing on CKS, both community support and enthusiasm may be enhanced. For example, a study by Butler et al. (2012) identified Turtles and Dugong as two traditional fishery resources in Torres Strait, Australia, which are the main source of livelihood for the local people. Over the years, the local people have accumulated a lot of traditional knowledge related to Turtles and Dugong. However, overfishing in recent years has drastically reduced the populations of these two species. In order to better protect the related resources, the local association absorbed the traditional knowledge of the local people and decentralized the governance to the community, which won the trust of the local people, promoted the formation of a co-management model and finally achieved the conservation goal.

3.4 Maintain the systematic function of AHS

3.4.1 Maintain the food and livelihood security of AHS

The biological resources on which CKS depend often play an important role in maintaining the food and livelihood security for local people. For AHS, cultivated plants and animals are usually closely related to local foods and customs, which are easily recognized as food CKS. On the one hand, they are the main sources of foods, providing a variety of foods and nutrition for local people. On the other hand, inedible plants and animals can be traded, so they are important means for the local people to maintain their traditional livelihood. A number of studies have already demonstrated this. For example, Freitas et al. (2020) point out that Arapaima, as a CKS in the Brazilian Amazon, is an important local commercial fishery resource. It not only provides food for the local people, but also supports the livelihood of local communities. When it was co-managed by the community as a CKS, the per capita income of Arapaima sales increased by 5-fold.

3.4.2 Conserve biodiversity in AHS

CKS can improve the preservation of local traditional varieties and maintain agro-biodiversity. First of all, it is necessary to identify and protect the material carrier of CKS, i.e., the biological resources such as species or varieties that have a significant impact on local society and culture. Objectively, this protects the biological resources of the AHS, helping to maintain local biodiversity.
Secondly, in general, some of the CKS in AHS are traditional varieties, which are important for the heritage diversity of organisms. Traditional varieties are more resistant to various adverse conditions. In the long-term development, they are gradually selected by farmers according to the local ecological environment and climatic conditions through hybridization and seed retention. Compared with modern varieties, traditional varieties can meet the needs of local people better and maintain stable yields in the absence of pesticides, fertilizers and other inputs of current technology. In addition, because most of the traditional varieties have better adaptability to the local natural environment, they have better resistance in the face of abiotic stresses such as droughts and floods (Zhang et al., 2016). These characteristics make traditional varieties rich in stable genetic traits. They are the reservoirs of germplasm resources in China, which are important for maintaining global agro-biodiversity.
Finally, both traditional knowledge and the interaction between biology and culture belong to the research category of CKS. The values contained in the traditional knowledge and culture are important for the conservation of local biodiversity and the ecosystem diversity. Therefore, the identification and conservation of CKS can be a useful tool for maintaining agro-biodiversity in AHS.

3.4.3 Promote the inheritance of traditional knowledge and technology in AHS

The traditional knowledge and culture contained in CKS are seen as a part of the local and traditional knowledge systems in AHS. The process of identifying and protecting CKS is also a process of discovering and protecting the traditional knowledge and technologies in AHS. For example, in the Congjiang Dong’s Rice-Fish-Duck System, there are some potential CKS which often appear in local village regulations and festivals, such as “Xianghenuo” and “Congjiang Local Ducks”. There are still some folk customs practiced by the Dong people. For instance, “San yue yue qing” means that on the eve of spring planting in March, the village leaders explain the rules and regulations on spring planting, fish and duck breeding, and mid-plowing management to everyone; and “Jiu yue yue huang” means that they focused on the regulations for harvesting, wintering of fish in the fields, and grazing of livestock in September. Through repeated preaching, these village regulations and treaties have become familiar to every villager. Among them, the cultivation, breeding, and harvesting techniques involved are all mastered, guiding the production and life order for generations. In turn, the above-mentioned traditional knowledge reinforces the community’s reliance on and identification with CKS. In Congjiang Dong’s Rice-Fish- Duck System, many prevailing festivals are held to record important agricultural events. When the festival comes, the local people must eat “Xianghenuo” rice, field fish and ducks, so that the unique local rice, fish, and duck varieties are protected. Thus, positive feedback between biology and culture is created, promoting the transmission of local traditional knowledge and technology systems and maintaining local species diversity.
In contrast, Berkes (2008), a famous ecologist, suggested that the extinction or decline of CKS would not only change the living conditions and spiritual beliefs of local people, but also bring negative impacts on the dissemination of local traditional ecological knowledge and technology. According to a survey conducted by the Yunnan Institute of Tropical Botany, Chinese Academy of Sciences, there were 800 traditional varieties of rice in Xishuangbanna in the 1970s. However, by the 1980s, more than 400 dry rice varieties were no longer being planted, and the knowledge related to these varieties was lost (Yunnan Institute of Tropical Botany, Chinese Academy of Sciences, 1982). Therefore, to protect CKS is to protect the traditional knowledge and technology associated with it.

3.4.4 Maintain the cultural values and social organizations of AHS

The cultivation and utilization of biological resources will have important impacts on the evolution of cultural values and social organization in a region. Li (2018) discussed how the structure of crop cultivation began to change with the introduction of a market economy system in a village in the Honghe Hani Rice Terraces System over the last 45 years. She reached three main conclusions. First, the main crops changed from rice and beans to cash crops such as vegetables, dendrobium and tea. Second, because of the change in planting structure, the pattern of land inheritance within family members has shifted, making the relationships between family members more tense than before. Thirdly, the confrontations between family relations triggered by political status disputes have increased, and the social network in the village has gradually shrunk from being based on kinship and neighbors to blood ties. The Indigenous Peoples International Declaration on Self-Determination and Sustainable Development believed that indigenous peoples’ cultural belief systems and worldviews are the basis for maintaining the sustainable development of local communities. Once local people start participating in the co-management of CKS, both local traditional knowledge and the subjective role of local communities will be enhanced, so that the local social organizations formed over a long time can also be maintained. Clans that rely on kinship, blood, and neighbor relationships will not be completely destroyed by the entry of modern social organizations such as governments and enterprises.

3.4.5 Maintain the landscape features of AHS

CKS play an important role in the formation and maintenance of AHS landscapes. Taking Honghe Hani Rice Terraces System as an example, alders (Alnus cremastogyne), which are closely related to local religious rituals, play an important role in maintaining the unique vertical landscape feature of the forest-village-terrace-water system, called “four elements isomorphism”. Generally speaking, the Hani terraces are located in cool mountainous areas with an altitude of over 2000 m. The dense forests are preserved there, which not only conserve water, but also provide abundant fruits and vegetables for the local people. In the sunny slopes at altitudes of 1400 to 2000 m, there are many villages, which are the place of daily life of the inhabitants. The semi-mountainous areas with altitudes of 600 to 2000 m are all covered with rice terraces, forming a water system with the characteristics of how high the mountain is and how high the water is. At the lowest point, there are rivers that receive the water flowing down from the terraces. This spatial pattern has the function of maintaining the water and soil, regulating the climate, and ensuring the safety of the village, as well as high aesthetic value. First of all, such a stable landscape has been created thanks to the water-supporting forest, the alder forest. Alder, as a unique tree species for Hani religious rituals, is usually treated as a sacred species and cannot be cut down at will. Secondly, rice cultivation plays an important role in the maintenance of the terraces, and crops such as soybeans and watercress are often grown on the field canals. These farming practices both prevent damage to the terraces and serve as a source of food for the local people. In addition, the structure of Hani’s houses is very unique. There are many mushroom houses made of bamboo, straw and adobe, divided into three levels. The first floor is a barn for raising animals, the second is for people and the third is for drying food and clothes. Not only are the building materials taken from various local creatures, but the building structure also has to consider the harmony of humans and animals, showing the wisdom of symbiosis between man and nature everywhere.
To sum up, a better understanding of the interactions between culture and organisms may be gained. When we focus on the identification and conservation of CKS, the biodiversity, traditional knowledge and cultural landscapes will be comprehensively protected. Eventually, the function and resilience of the AHS will be improved as a whole (Fig. 2).
Fig. 2 Conservation of AHS function by cultural keystone species

4 Discussion

For AHS, there are still many challenges in the identification and application of CKS. In the future, the research of CKS can be strengthened in the following two areas.
First, the awareness of the importance of CKS needs to be raised. There is still relatively little discussion of CKS in China, with most studies focusing on the interactions between organisms and minorities. For example, Luo et al. (2019) investigated edible wild plants in the Honghe Hani Rice Terraces System and found that most of the plants had great economic potential. The edible wild plants can help the local people recognize the relationships and value between wild species and related traditional knowledge. Zhang (2019) found that the traditional cultivation and utilization of taro played an important role in the production practices of Native Hawaiians. To the locals, taro is not only a food, but also a sacred object that is inextricably linked to their ancestors. However, none of the studies mentioned the concept of CKS, which indicates that this research perspective is thought to have still not attracted the attention of relevant scholars. In the research of AHS, Ding (2021) studied the identification and conservation of CKS in AHS. But this study mainly focused on the enumeration of CKS of the case sites, lacking any revelation of the interactions between biology and culture. In the future, we should pay more attention to the theoretical research of CKS. Researchers should analyze the impact of CKS on the function of AHS and use it for management practices.
Second, to use the concept of CKS to promote community participation in the co-management of local resources and ecological conservation, it is critical to identify the CKS in a given area. Using quantitative research methods can help to quickly compare the importance of different species in the living practices of local people and to integrate the locals’ opinions of them. However, there is no uniform quantitative standard for the study of CKS identification. When Garibaldi and Turner (2004) proposed the CKS concept, the index of the identified cultural influence of CKS was designed to measure the cultural importance of species. This index includes six criteria: intensity, type and multiplicity of use; naming and terminology in a language; roles in narratives, ceremonies, or symbolism; persistence and memory of use in relationship to cultural change; level of unique position in culture; and extent to which it provides opportunities for resources. Although this index includes different aspects of culture and provides a brief framework for identifying CKS, in the actual process of identifying CKS, the indicators are determined based on the subjective perceptions of researchers or respondents. Therefore, the results obtained are always restricted by the subjectivity of the researchers (Davidson et al., 2012). In addition, ethnobotanists often use cultural significance index (Reyes-García et al., 2006), cultural value index (Da Silva et al., 2006), use-value (UV) and the frequency of utilization index (FUI) (Luo et al., 2019) to identify CKS. However, the results identified by these methods are difficult to replicate and validate (Coe and Gaoue, 2020), and there is a lack of measurements of human-biological interactions and a deeper understanding of the culture. The identification of CKS involves the influence of cultural factors related to local communities, including the religious beliefs, values, and cosmology of the local people. Determining how to measure the importance of cultural factors is an important issue to be resolved in the future.

5 Conclusions

As a composite of the organisms and cultures that have shaped the cultural identity of a people, CKS contains complex interactions between organisms and cultures. It can promote the comprehensive conservation of biodiversity and cultural diversity with the advantages of easy management and high community acceptance, contributing to the resilience of the socio-ecological systems. The identification of CKS is a breaking point in promoting the conservation of AHS, which are typical SES. First, CKS help to quickly identify the key elements of AHS and prioritize the conservation objects. Second, because CKS are easily recognized and supported by local people, they can promote community participation in the conservation and development of the AHS. In addition, the identification of CKS has a significant role in food and livelihood security, biodiversity conservation, traditional knowledge and technology transmission, social organization maintenance, and cultural landscape maintenance in the AHS, which helps to achieve the conservation goals of the AHS.
Agnoletti M, Rotherham I D. 2015. Landscape and biocultural diversity. Biodiversity and Conservation, 24(13): 3155-3165.


Ann G. 2009. Moving from model to application: Cultural keystone species and reclamation in fort McKay, Alberta. Journal of Ethnobiology, 29(2): 323-338.


Berkes F. 2008. Sacred ecology (2nd ed.). New York, USA: Taylor & Francis Group.

Butler J R A, Tawake A, Skewes T, et al. 2012. Integrating traditional ecological knowledge and fisheries management in the Torres Strait, Australia: The catalytic role of turtles and dugong as cultural keystone species. Ecology and Society, 17(4): 34-52.

Chen B, Akamine H. 2021. Distribution and utilization of homestead windbreak Fukugi (Garcinia subelliptica Merr.) trees: An ethnobotanical approach. Journal of Ethnobiology and Ethnomedicine, 17: 11. DOI: 10.1186/s13002-021-00434-3.


Coe M A, Gaoue O G. 2020. Cultural keystone species revisited: Are we asking the right questions? Journal of Ethnobiology and Ethnomedicine, 16: 70. DOI: 10.1186/s13002-020-00422-z.


Cristancho S, Vining J. 2004. Culturally defined keystone species. Human Ecology Review, 11(2): 153-164.

Da Silva V A, Laise D H C A, De Albuquerque U P. 2006. Revising the cultural significance index: The case of the Fulni-ô in northeastern Brazil. Field Methods, 18(1): 98-108.


Davidson-Hunt I J, Turner K L, Mead A T P, et al. 2012. Biocultural design: A new conceptual framework for sustainable development in rural indigenous and local communities. Paris, France: Institute Veolia Environnement Books.

Ding L B. 2021. Study of cultural keystone species and their protection in agricultural heritage sites-The case study in Honghe County, Yunnan and Aruhorgin, Inner Mongolia. Diss., Beijing, China: University of Chinese Academy of Sciences. (in Chinese)

Ellen R. 2006. Local knowledge and management of sago palm (Metroxylon sagu Rottboell) diversity in south central Seram, Maluku, eastern Indonesia. Journal of Ethnobiology, 26(2): 258-298.


FAO. 2002. Report of the FAO workshop of potential stakeholders and steering committee on Globally-Important Ingenious Agricultural Heritage Systems. Rome, Italy: FAO Land and Water Division.

Freitas C T, Lopes P F M, Campos-Silva J V, et al. 2020. Co-management of culturally important species: A tool to promote biodiversity conservation and human well-being. People and Nature, 2(1): 61-81.


Gaoue O G, Coe M A, Bond M, et al. 2017. Theories and major hypotheses in ethnobotany. Economic Botany, 71(3): 269-287.


Garibaldi A, Turner N. 2004. Cultural keystone species: Implications for ecological conservation and restoration. Ecology and Society, 9(3): 1.

He L, Song Y J, Long C L. 2020. Genetic diversity of rice landraces in Yuanyang Hani Terraced Rice Fields under in-situ conservation: Research progress. Chinese Agricultural Science Bulletin, 36(10): 87-94. (in Chinese)

Jacques-Coper A G, Cubillos G, Ibarra J T. 2019. The Andean condor as bird, authority, and devil: An empirical assessment of the biocultural keystone species concept in the high Andes of Chile. Ecology and Society, 24(2): 35. DOI: 10.5751/ES-10939-240235.


Jiao W J, Cui W C, Min Q W, et al. 2021. A review of research on Agricultural Heritage Systems and their conservation. Resources Science, 43(4): 823-837. (in Chinese)

Li H Y. 2021. Identification and management of key elements of the Agricultural Heritage Systems: Taking terraced systems as examples. Diss., Beijing, China: University of Chinese Academy of Sciences. (in Chinese)

Li W H. 2015. Agri-cultural Heritage research and conservation practices: Progress and perspectives. Journal of Agro-Environment Science, 34(1): 1-6. (in Chinese)

Li W Q. 2018. Crops, market and social relations: A case study of Hani in a village. Journal of Original Ecological National Culture, 10(4): 98-105. (in Chinese)

Long C L, Xue D Y, Feng J Z. 2009. Traditional management of natural resources in ethnic areas. Beijing, China: China Environmental Science Press. (in Chinese)

Luo B S, Liu B, Zhang H Z, et al. 2019. Wild edible plants collected by Hani from terraced rice paddy agroecosystem in Honghe Prefecture, Yunnan, China. Journal of Ethnobiology and Ethnomedicine, 15: 56. DOI: 10.1186/s13002-019-0336-x.


Mao S X, Shen Y, Deng H B. 2017. Progress in biocultural diversity research. Acta Ecologica Sinica, 37(24): 8179-8186. (in Chinese)

Maxwell K H, Ngāti Horomoana T W H, Arnold R, et al. 2018. Fishing for the cultural value of kahawai (Arripis trutta) at the Mōtū River, New Zealand. New Zealand Journal of Marine and Freshwater Research, 52(4): 557-576.


McMillen H L. 2008. Conserving the roots of trade: Local ecological knowledge of ethnomedicines from Tanga, Tanzania markets. Diss., Honolulu, USA: University of Hawaii.

Min Q W. 2020. Research priorities, problems and countermeasures of Important Agricultural Heritage Systems and their conservation. Chinese Journal of Eco-Agriculture, 28(9): 1285-1293. (in Chinese)

Min Q W. 2021. Agri-cultural heritage: An interdisciplinary field with development prospects. Journal of Resources and Ecology, 12(4): 437-443.

Paine R T. 1969. A note on trophic complexity and community stability. The American Naturalist, 103(929): 91-93.


Parviz K. 2009. Conservation and adaptive management of Globally Important Agricultural Heritage System (GIAHS). Resources Science, 31(1): 4-9. (in Chinese)

Pei S J. 2011a. General introduction to three-decades development of ethnobotany and perspectives in China. Journal of Minzu University of China (Natural Sciences), 20(2): 5-9. (in Chinese)

Pei S J. 2011b. Traditional culture and biodiversity conservation. Bulletin of Chinese Academy of Sciences, 26(2): 190-196. (in Chinese)

Pilgrim S, Pretty J, Adams B, et al. 2009. The intersections of biological diversity and cultural diversity: Towards integration. Conservation and Society, 7(2): 100-112.


Platten S, Henfrey T. 2009. The cultural keystone concept: Insights from ecological anthropology. Human Ecology, 37(4): 491-500.


Reyes-García V, Huanca T, Vadez V, et al. 2006. Cultural, practical, and economic value of wild plants: A quantitative study in the Bolivian Amazon. Economic Botany, 60(1): 62-74.


Singh R K, Garnett S T, Zander K K, et al. 2015. Paisang (quercus griffithii): A keystone tree species in sustainable agroecosystem management and livelihoods in arunachal pradesh, India. Environmental Management, 55(1): 187-204.


UNESCO. 1988. Declaration of Belem. Third UNESCO science and culture forum. Towards eco-ethics: Alternative visions of culture, science, technology and nature. Belem, Brazil: 6-10.

Winter K, McClatchey W. 2009. The quantum co-evolution unit: An example of Awa (Kava-Piper methysticum G. Foster) in Hawaiian culture. Economic Botany, 63(4): 353-362.


Yang L. 2019. Research on farmer’s livelihood transformation in GIAHS site-A case study of Zhagana Agriculture-Forestry-Animal Husbandsry Composite System. Diss., Beijing, China: University of Chinese Academy of Sciences. (in Chinese)

Yang Y H, Bai K Y, Devra J, et al. 2019. Xishuangbanna cucumber landraces and associated traditional knowledge. Biodiversity Science, 27(7): 743-748.


Yunnan Institute of Tropical Botany, Chinese Academy of Sciences. 1982. Collection of reports on tropical plant research papers. Kunming, China: Yunnan People’s Publishing House.

Zhang D, Min Q W, He L, et al. 2016. Agrobiodiversity features, conservation and utilization of China’s Globally Important Agricultural Heritage Systems. Chinese Journal of Eco-Agriculture, 24(4): 451-459. (in Chinese)

Zhang Y. 2019. Taro planting and Hawaiian ethnic identity. Journal of Original Ecological National Culture, 11(6): 1-8. (in Chinese)