A framework to categorise the cultural significance of freshwater fauna

A framework to categorise the cultural significance of freshwater fauna | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article A framework to categorise the cultural significance of freshwater fauna Phoebe Griffith, Sonja Jähnig, Rebecca Tharme, Fortunate Phaka, and 9 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6830897/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Freshwater ecosystems support rich biological and cultural diversity, each enhanced through biocultural co-evolution. However freshwater species, and their relationships with people, face escalating pressures from global change, threatening ecosystem function and human wellbeing. Addressing this crisis effectively and ethically requires transdisciplinary approaches that recognise the complex, culturally embedded interactions between people and freshwater fauna. Existing frameworks for integrating the cultural and ecological dimensions of freshwater fauna are either too general or conceptually misaligned to support practical application, and are often based on binaries such as ‘tangible and intangible’ that artificially segregate meaning and practice. To bridge this gap, we developed a novel framework for systematically categorising the cultural significance of freshwater fauna, drawing on pre-existing frameworks and human-freshwater-fauna interactions (n = 612) from interdisciplinary literature. The resulting framework comprises seven overarching domains – Consumptive Use; Cultural Landscapes, Sites and Stewardship; Knowledge Systems; Worldviews, Beliefs and Identities; Cultural and Social Practices; Recreation, Leisure and Tourism; and Creative Expression – each subdivided into defined categories and subcategories. Applicable across diverse cultural, taxonomic and geographic contexts, our framework provides a pragmatic tool to support more holistic and inclusive approaches to freshwater ecosystem research, management, governance, and cross-cultural cooperation. Scientific community and society/Water resources Earth and environmental sciences/Environmental social sciences/Environmental impact Scientific community and society/Social sciences/Interdisciplinary studies Figures Figure 1 Figure 2 Figure 3 Introduction Inland waters have played an essential role in human societies since time immemorial, with freshwaters – including rivers, lakes and wetlands – shaping the history of humanity 1 , 2 . Even today, 90% of the world’s population lives within 10 km of surface freshwater bodies 3 . The disproportionately high biodiversity of freshwaters, relative to their small proportion of the earth's surface, is well established 4 . Freshwater ecosystems are also known for their formative and critical roles in the lifeways of peoples of different cultures 5 – 7 . Consequently, freshwaters have high biocultural diversity and value. Biocultural diversity explicitly recognises the deeply intertwined, co-evolved and dynamic relationships between biological and cultural diversity, and the often place-based relationships that underpin socio-ecological systems 8 , 9 . Nonetheless, both biological and cultural diversity are imperilled. One quarter of freshwater fauna species are threatened with extinction 10 , with the potential impacts of this loss on the human cultures and communities they interact with extending far beyond a count of threatened species. Cultural diversity is also at risk, with nearly half of the world’s spoken languages on the brink of extinction 11 . The loss of these languages is closely tied to both cultural and biodiversity declines 12 . Simultaneously, erosion of relationships between people and freshwater biodiversity both exacerbates species extinction risks and negatively impacts human wellbeing 13 , 14 , particularly through the loss of Indigenous and other local knowledges 15 . As much of the biodiversity that exists today has co-evolved with human cultural diversity, in many contexts their maintenance is inseparable 12 , 16 . For biodiversity conservation, management and governance to be ethical, equitable and effective, therefore, cultural context should be a central consideration 17 – 19 . Such consideration is likely to be particularly key in freshwater environments, where humans and nature are deeply entwined 20 . However, the vital role of culture in conserving biodiversity – and of biodiversity in sustaining cultures – has yet to be fully acknowledged in the global discourse on environmental protection 21 . In fact, while the multi-directional relationships and nature-culture nexus are well conceptualised in some worldviews, particularly those of many Indigenous Peoples 22 – 24 , the complex and culturally informed relationships between people and other elements of nature have been largely overlooked in the Western scientific worldview. Consequently, until recently, relations between nature and culture have been largely absent in international and national policy instruments addressing the biodiversity crisis 25 – 27 . This situation is changing, however as reflected in the Convention on Biological Diversity Decision 15/22 on Nature and Culture 28 and the adoption of the concept of nature’s contributions to people (NCP) by the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES). The IPBES defines NCP as “all the contributions, both positive and negative, of living nature to people’s quality of life” (Díaz et al., 2018, p. 270), recognising the critical role of culture in defining all relationships between people and other nature. Such changes are important, as many sustainable solutions to environmental degradation are found at the intersection of nature and culture, in the form of reciprocal relationships with other-than-human nature 29 . Conceptual and applied frameworks aid in understanding and describing the types of diverse relationships that exist between people and freshwater biodiversity. Such frameworks help communicate the context-specific cultural interactions between people and nature to those in other communities, aiding in global uptake and understanding of biocultural approaches 9 . For example, the Ramsar Convention, a critical policy instrument for freshwater ecosystems, has recognized the importance of culture since the treaty was adopted in 1971 6 , and has set out a typology of significant wetland-related activities to systematise the types of cultural significance wetlands, as places, hold for people 30 . Using a different lens, Lynch et al. (2023) applied the IPBES NCP reporting categories to identify various contributions of freshwater biodiversity to people, including non-material cultural contributions. However, in their application, ‘culture’ is treated as a singular undifferentiated category, separate from other culturally rich categories, such as food or recreation, as is also the approach in the Ecosystem Services framework 20 . Interpretation of the contribution of freshwater species thus seems to depend on the concept of culture as considered by those applying the framework, with potential for a narrow or biased scope. While these kinds of frameworks have progressed, we still lack a robust, unifying framework for categorising and systematically understanding, including for practical application, the importance that animal species found in freshwaters may hold for different communities and cultural groups. The absence of a such framework for assessing the cultural significance of freshwater species is particularly problematic in that species remain a primary unit for biodiversity conceptualisation and practical conservation action 31 – 33 . Biocultural approaches that consider the place-based interrelationships of people and nature at the species level are useful for efforts in transdisciplinary conservation 9 . Consequently, a framework for connecting the scientific concepts of species (or other taxonomic groups or taxa) to their social and cultural contexts can be a pragmatic tool for knowledge sharing across disciplines and among cultures. Such a framework could be useful, for example, to communicate about migratory species that traverse multiple human communities, territories, and cultures. The conservation of such species and associated cultural diversity requires cooperation across these diverse groups. To illustrate, freshwater eels ( Anguilla species, spp.) are one such taxonomic group that migrates vast distances between fresh waters and oceanic breeding grounds to complete their life cycle 34 . The giant mottled eel ( A. marmorata ) is crucial to communities across the Pacific, including in Melanesia, Polynesia and Kiribati 35 – 38 . The species is an ecological indicator, and holds spiritual and economic significance 37 , 39 . In some areas, this species symbolizes ancestral connections, while in others, it is a totem species and or vital food source. A framework that allows regional communication and cooperation in managing these migrant populations would therefore be aided by mutual understanding of this diversity in human-eel relationships. In this context, we developed a first of its kind framework for capturing human-freshwater fauna interactions across diverse knowledge systems and disciplines, drawing on a broad and interdisciplinary literature. By categorising these relationships, we can conceptualise what cultural significance of species looks like practice. We define ‘cultural significance’ as the importance of a particular taxon’s role within a specific culture 40 , 41 , including the values, importance, knowledge, meaning, and relationship of that species to a particular cultural group 42 . By proposing a framework to categorise different culturally informed interactions between people and freshwater fauna, we hope to provide a pragmatic way to improve understanding and strengthen the integration of cultural significance in safeguarding of natural and cultural diversity and heritage. Methods Framework Development and Category Definitions To establish a unique framework to address the cultural significance of freshwater fauna, we first examined the ways in which elements of culture have already been framed and grouped, including for freshwater ecosystems. We developed the initial draft categories for our framework in large part through reviewing the categorisations used in five frameworks known to the authors: (i) the Ramsar Typology of Significant Wetland-Related Activities 30 , (ii) reporting categories for Nature’s Contributions to People 25 , (iii) UNESCO Intangible Cultural Heritage domains 43 , (iv) the International Union for Conservation of Nature (IUCN) Species Use Database 44 , and the (v) South African ethnoherpetological elements of culture 45 . We selected these frameworks either for their broad relevance to human interactions with freshwater species, and/or for their in-depth consideration of key aspects of cultural and use dimensions. We also chose them for their thematic diversity, established or emerging application in policy and practice, and/or finer-scale delineation of certain types of human-fauna relationships. To create an integrated and robust framework, categories and domains from each framework were compiled, reviewed for conceptual alignment, and consolidated or adapted, where duplicate or overlapping classifications existed. For example, some categories we directly drew from past frameworks, such as categories ‘social practices, rituals and festive events’ and ‘performing arts’ from the UNESCO domains of intangible cultural heritage. In other cases, we consolidated categories, for example, ‘food and feed’ (NCP reporting categories), ‘food processing’ (Ramsar Typology) and ‘gastronomy’ (ethnoherpetological elements) to the category in our framework of ‘food and culinary heritage’. Where additional differentiation was feasible and logical, categories were further subdivided into finer subcategories to, as closely as possible, group together culturally informed interactions between people and freshwater animals, systematically describing similarities in the ways different cultural groups relate to species. Defining culture, cultural groups, and species We focus on biocultural diversity, defined as “the range of human values and practices related to the dimensions of biodiversity at landscape, species and genetic levels” 8 , encompassing both biological and cultural diversity, and the interactions between the two that shape mutual coexistence. Our scope includes all peoples and is not limited to rural areas, as is sometimes the case in biocultural studies. We follow Reyes-Garcia et al. (2023) 9 , who define a cultural group as a community of practice who share a core set of beliefs, patterns of behaviour, and values. We identified cultural groups to the most specific level possible, based on literature provided, for example, ‘Tharu People from Chitwan district, Nepal’ rather than ‘Indigenous People, Nepal’ or ‘Tharu People’. Where reports from the literature provided only broad descriptors of sociocultural identity (e.g. Native Americans), we used these descriptors. We used the level of ‘species’ (using scientific taxonomic nomenclature) as the unit of analysis for the biological diversity component in our framework, as this is the most widely used and recognised unit in conservation science and management 33 , for instance, the IUCN Red List of Threatened Species 46 . People often classify wildlife using folk taxonomies, which may align but are not always synonymous with scientific taxonomy. In our approach, we tie the cultural roles of a scientific species to a particular ‘folk taxon’ or ‘ethnospecies’ 47 – 49 which required matching biological species with ethnospecies. In most instances, this was straightforward, as the source provided the scientific name of the species, the corresponding folk or vernacular name, or an image of the species. For example, ‘pike’ in the United Kingdom corresponds to Esox lucius , and images of a pike cannot be mistaken for those of any other local species. Where we found a lack of alignment, due to a folk-taxon corresponding to more than one scientifically defined species, all potential corresponding species, or an entire genus or family, were listed as the scientific species. For example, in Europe, multiple languages commonly use a single name - such as sturgeon (English), Stör (German) or есетра (Bulgarian) – to refer to all sturgeon species, especially Acipenser spp. and sometimes also Huso huso. In another example, in popular culture, ‘crocodiles’ is used to refer indiscriminately to all true crocodile species of the family Crocodylidae. Where there is a lack of information in the literature to differentiate between candidate species that may correspond to a folk taxon, all potential species were noted, along with the uncertainty in attribution. In some instances, we only included the folk or vernacular name, due to an overwhelming number of corresponding species. For example, for some of the rituals related to frogs there were well over 50 potential species to include. In these instances, we included a note to explain this. Example Collection With a draft complete framework designed, we then iteratively tested out and refined how well the initial categories fared against a first set of collated examples that covered the full range of categories defined and major world regions. We collected case examples of relationships between humans and freshwater fauna that were portrayed as or could be interpreted as cultural. Relationships between humans and freshwater fauna included knowledge, practices, beliefs and uses. Examples were collated from peer-reviewed publications, grey literature, personal knowledge of the authors and other disciplinary specialists, field data from authors’ own work, online museum collections, as well as through a snowballing approach (i.e., searching relevant references cited in the studies identified). For each example, we collected the most specific information available in the reference or record referring to species, sociocultural and/or ethnolinguistic groups, and a summary of the relationships and knowledge (Table 1 ). We aimed to include a geographically, culturally, and taxonomically diverse set of examples and, therefore, actively sought out examples from continents where we initially lacked examples. We included examples from both native and alien species. We aimed for a minimum of three examples for each initial category (and at least one for a subcategory). With expansion of the dataset during adaptive testing of the framework (see below), we collected some 612 case examples on which we based the finalisation of the framework. As the focus of this study is outlining the framework, we then selected the most informative subset of these cases (n = 232) to illustrate the framework for this paper, comprising a first knowledge base on which to continue to build (Supplementary Table S3). Many of the examples represent secondary sources of knowledge. A potential limitation in such instances is that different actors use different lenses to interpret and report knowledge. Wherever possible, therefore, we prioritised the sources closest to the original context. For instance, where multiple sources were available, we would favour those produced by the communities themselves, such as drawing on cosmogonies from official websites of First Nations or Aboriginal groups, rather than relying on colonial accounts. Table 1 Information collected for each example taxon of cultural significance Taxonomic Information Information on Sociocultural Group Information on the nature of the relationship and the associated knowledge • Family • Genus • Species • Name or description of sociocultural group (e.g. ethnolinguistic group, societal group) • Place of knowledge/relationship • Country (or countries) of knowledge/relationship • Folk name(s) of ethnospecies used by give sociocultural group • Description or illustration of the relationship and/or knowledge • Notes on how ethnospecies was related to scientific species (if not immediately clear and intuitive e.g. if there is under differentiation of species within a folk taxon) Assigning case examples to domains, categories and subcategories A specific definition was developed for each category and subcategory in the framework. Where possible, established definitions were drawn from the literature. For example, the category “Tourism” uses the definition of tourism from the United Nations Statistical Commission (2008) 50 . In most cases, a specific definition was developed, drawing on broader conceptualizations found in the existing literature. For example, the category ‘Worldviews and belief systems’ is defined as “the species plays a key or specific role in a cultural worldview, which is the perceptual framework of a cultural group, in which worldviews are ‘socially constructed realities which humans use to frame perception and experience’ and refers to the ‘psychological, cognitive and affective determinants of behaviour’ 51 . It includes beliefs, such as on cosmology, cosmogonies, the afterlife, goals, values, and ethics. Cultural worldviews may or may not be distinguishable from religious or spiritual beliefs, depending on the culture 52 .” Where the definition described a comparatively straightforward concept, we did not reference other literature as, for example, for the category “Representation in Music” which is defined as “the species is represented in and/or inspiration for the creation of music, such as songs, rhymes, chants, and instrumental music.” We assigned each example in our dataset to one or more categories or subcategories in the draft framework. Where examples did not fit comfortably within the framework structure, (sub)categories were adjusted or new (sub)categories were developed, through an iterative process. Where examples illustrated multiple facets of the relationship, each of the different facets were included in the different relevant categories and, where necessary, in multiple categories to ensure full representation of cultural significance. Framework refinement continued until all collected examples could be readily assigned to one or more categories. Through this process, we refined the categorization structure to ensure theoretical relevance for freshwater fauna, with reassessment of the specificity and distinctiveness of each category ensuring that the framework was both inclusive and analytically robust. Finally, we organised categories into groupings of thematically similar relationships, comprising seven overarching domains, which were named to reflect their internal categories (Fig. 1 ). Domains are thematically defined by the groupings of categories attributed to them. For example, the domain (7) thematically defined as ‘creative expression’, is made up of the individually defined categories of visual arts (7.1), performing arts (7.2), language arts (7.3) and other forms of creative inspiration (7.4). The resultant framework was a hierarchical three-tier system, with broad domain subdivided into categories, which were divided into subcategories where necessary (Fig. 1 , Table S1 ). The hierarchical organization was designed to capture increasing specificity in the nature of human-species relationships, while maintaining conceptual clarity and practical applicability across diverse cultural contexts. Our example dataset includes multiple examples for each domain, category and subcategory, from which we have selected those that best illustrate each of these framework elements. We also included some specific cultural group and species pairings in the results repeatedly, to illustrate how the relationship between a given species and culture can have multiple facets. Similarly, we included relationships of different cultures with the same species or taxon, to illustrate how one species can have multiple significance across multiple cultures. Positionality statement This research aims to provide a framework for identifying and categorising types of human-freshwater fauna relationships across multiple cultures, with the goal of improving cross-cultural comparisons and understanding of these relationships. Our author team includes people from different nationalities, disciplinary backgrounds, and positionalities, and both Indigenous and non-Indigenous scholars. We acknowledge that our approach, and the framework construct itself, is largely influenced by Western epistemic traditions, and is driven by the aim to facilitate understanding of the biocultural importance of freshwater fauna to scholars, practitioners, policymakers, stakeholders and rightsholders. We hope to contribute to respectful, clearer cross-cultural communication and recognition of cultural significance as an integral and vital part of freshwater ecosystem management and conservation. We appreciate that Indigenous Peoples, and different cultural groups or communities, might conceptualise, categorise, and understand relationships very differently, and through different lenses these categories may not be universally applicable, meaningful, or appropriate. We recognise these limitations of the framework, as well as those considered in the below and offer it, not as a universal approach, but as one possible tool to build a coherent evidence base that supports cross-cultural dialogue across knowledge systems. With this framework, we seek to provide a pragmatic and respectful tool for recognising the plurality of human-freshwater fauna connections, and for making these relationships more visible and actionable in conservation and heritage contexts. Considerations on use and application of the framework A comprehensive understanding of the cultural significance of species necessitates deep engagement with the complex and often nuanced cultural contexts in which human–other-than-human relationships are embedded. These relationships are inherently context-specific and can be challenging to convey to those outside a cultural group 53 . The framework we propose offers a structured approach for exploring cultural significance. It is not intended as a substitute for participatory, inclusive methodologies that respect the rights and autonomy of cultural groups, particularly in conservation and related applications. It serves as a tool to aid in systematising knowledge, and as a starting point for exploring the diverse ways in which people relate to freshwater species. Similarly, assignment of categories can be challenging to those without a deep understanding of a culture. We sought to reduce this limitation as much as possible by dividing categories at the most easily ‘quantifiable’ part of the relationship (i.e., the human-animal or human-representation interaction), rather than using a framework based on values. For example, recreational fishing may be valued for a range of culturally specific reasons such as fun, additional food, or relationship-building. However, we class it as recreational fishing based on the activity, rather than the underlying justification or values. As with all research involving people, there are important ethical considerations in the collation and mobilisation of data on the relationships between human and other elements of nature 54 . A careful, sensitive and respectful approach to knowledge is particularly important with Indigenous and other local community knowledge, and we highlight the importance of proper engagement in understanding cultural engagements with freshwater taxa 55 . For example, some knowledge and other aspects of relationships between species and a given culture are sacred and/or secret knowledge 56 , and maintaining the sacred or secret nature of such knowledge is paramount. Our framework provides one approach to discuss or reference a relationship without giving any specifics on the knowledge itself, with the categories providing a potential placeholder for such knowledge in databases and/or discussions. This needs to be done on a case-by-case basis, as in some circumstances, collection or categorisation of knowledge could be unethical or inappropriate. We urge all users of this framework to consider which data are included in a cautious, respectful and rights-based manner. Results The framework (Fig. 1 , Table S1 ) organises culturally significant relationships between people and freshwater fauna into seven overarching domains, representing thematic groupings which broadly capture the diversity of human-fauna interactions. Each domain is divided into multiple categories, and some categories are further subdivided into subcategories. The full set of categorized relationships is provided in Fig. 1 . To facilitate interpretation, detailed definitions of each category and subcategory are provided in the Supporting Information (Table S2). To explain the framework, we use a selection of brief references to examples, all of which are further detailed in the Supporting Information (Table S3), alongside a larger set of explanatory examples (n = 232). Domain 1: Consumptive Use This domain encompasses human cultural interactions involving the consumptive use of the species, alive or dead, and related knowledge. A particularly knowledge rich theme, it includes most interactions typically considered ‘material’, ‘tangible’, or ‘provisioning’ —such as food, material, and medicinal uses 20 — as well as ‘intangible’ or ‘immaterial’ aspects of such uses. For instance, the fish Tenualosa ilisha is widely eaten in South Asia and exemplifies multiple categories of this Domain. In Bangladesh, it is important for food security (1.1.1) and nutritional security (1.1.2), accounting for over 12% of Bangladesh’s total fish production and 1% of national gross domestic product 57 . Called Ilish in Bengali, it is consumed in ritual and social practices (1.1.4), such as Bengali New Year. Bengali Hindus also offer the fish to deities in other festivals 58 . In Sindh, Pakistan, it is a prized food called Palla , which is offered to guests as a sign of honour and respect (1.1.3 and 1.1.4) (Fig. 2 ). Consumptive relationships can also have negative effects, as is the case of the fish Aymara ( Hoplias aimara ) of French Guiana. While its consumption is a delicacy (1.1.3) for many local communities (Griffith & Mangione, unpub. data), eating the fish poses a health risk due to mercury contamination 59 . The species complex collectively known as the Chinese Giant Salamander ( Andrias spp.) is also a delicacy , as well as status symbol food in mainstream urban Chinese culture 60 . The latter cultural connection, along with local consumption in communities across its range, has pushed the salamander close to extinction in the wild 60 . The category knowledge related to capture (1.2) includes traditional practices that enable locating, capturing, and killing of a species. Tharu fishers in Chitwan Nepal employ a method of attracting eels by constructing underwater stone piles (Griffith, unpub. data), represented under the subcategory of tracking, locating and attracting knowledge (1.2.1). The knowledge of how to weave Māori eel traps (Hīnaki) to capture Anguilla spp. or tuna in the Māori language, is capturing knowledge (1.2.2) 61 . Healing and pharmacological uses also form a category within Consumptive Use . Many species are used in traditional healing systems (1.3.1). For example, red toads ( Schismaderma carens ) are sold in traditional medicine markets of Johannesburg, South Africa 62 . Similarly, fisher families of the Tocantins River, Brazil, use the fat of the poraquê eletrico (electric eel, Electrophorus electricus ) as medicine and the electric shocks to treat rheumatism 63 . Certain species are used in conventional medicine and research (1.3.2), like zebrafish ( Danio rerio ), a prominent model organism for the international scientific community 64 . Other pharmacological uses (1.1.3) include poisons, mind-altering drugs, and drugs for spiritual or cultural rituals. For example, AmaZulu and AmaXhosa peoples of South Africa use nwenya, Nile crocodile ( Crocodylus niloticus ), to make poisons and potions for strength 65 , while Matsés hunters, Peru, use the Sapo (derived from secretions of the frog dowkiet, Phyllomedusa bicolor) to enhance hunting success 66 . C onsumptive use also includes a category that captures the use of the animal or its parts in crafts, artefacts or tools (1.4.1), or items to be worn, such as clothing or jewellery (1.4.2). Freshwater pearls from the cockscomb pearl mussel, 褶纹冠蚌 ( Cristaria plicata ), have been used for millennia in China, for ornamentation (1.4.2) 67 . In Medieval Western Europe, isinglass from sturgeon ( Acipenser spp. and Huso huso) swim bladders was used as glue in book illumination 68 . Other categories include use for energy (1.5), such as oil from the eggs of the Arrau turtle ( Podocnemis expansa , Tartaruga) that Portuguese settlers used for lighting 69 . The name ‘candlefish’ attributed to the Eulachon Thaeleichthy pacificus refers to the traditional practice of using the dried fish as a candle (1.5.1, energy from primary use ), a method sometimes practiced by the Nisda’a First Nations people in Canada. They call the fish halmotkw, which can be translated as ‘salvation fish’, as the anadromous run of the fish was important for food security (1.1.1), and today remains important for consumption in ritual or social practices (1.1.4), especially the grease of the fish (1.1.6, food processing ) 70 . The use for companionship and decoration (1.6) is also a category of this domain: 龙鱼, the Asian arowana ( Scleopages formosus ) is an exceptionally popular aquaria pet in Chinese cultures 71 . Finally, non-use removal (1.8) refers to cases where a species is killed or removed without any intention of use, often due to a perceived negative cultural value, such as bad luck, as an ill omen, or belief in a negative interaction with another preferred species. For example, alligator gar ( Atractosteus spatula ) have been viewed in recreational fishing communities in the USA as detrimental to ‘game fishes’ and have been persecuted to the point of eradication in some areas 72 . Domain 2: Cultural Landscapes, Sites and Stewardship This domain captures how the relationship between people and certain freshwater species shapes landscapes, including through the emergence of cultural landscapes, sites and places of settlement or worship, and stewardship practices (Fig. 1 ). Sites and structures (2.1) refers to locations shaped through connections with specific species. We identify three time periods that indicate the origin of the sites and structures in their current (or final) location, aligned with the Ramsar typology of wetland-related activities 30 : ancient and prehistoric (pre-500CE); post-classical (500-1599CE); and traditional and modern (post-1599CE). Ancient and pre-historic sites and structures (2.1) include Mount Sandel, a Mesolithic Irish settlement site, likely sited due to reliance on eel, eascann ( Anguilla anguilla ), and salmon, bradán ( Salmo salar) , from the River Bann 73 . Small-scale Karpfen (common carp, Cyprinius carpio ) farming pond landscapes in Bavaria, Germany, have been maintained since the Middle Ages, and therefore fall under the subcategory of a post-classical site (2.2). They are listed as Intangible Cultural Heritage by the German Commission for UNESCO (2023). The Chabdhi Barahi Temple in Tanahun, Nepal, established by the local Magar community in 1939, on instruction of the Goddess Barahi Devi, is a traditional and modern site and structure (2.3). Sacred Rajabaam eels ( Anguilla bengalensis ) in the temple pools are a key agent in worship (Fig. 2 ), viewed variously as an incarnation, messenger, or protector of the goddess (Griffith & Basnet unpub. data). It is impermissible to fish for these eels locally. In Chile, a traditional and modern site and structure (2.3) has arisen in Lake Llanquihue. In the 1970s, the industry based on non-native salmonids created a new ecosystem for the purpose of aquaculture and recreational fishing, now a multi-million dollar industry 74 , although the resultant declines in native species lead to conflicting views on the emergence of this new and novel ecosystem dominated by alien species 75 . Stewardship practices (2.2) are actions taken to protect, care for, or responsibly use the environment 76 , and reflect the cultural aspects of peoples’ relationships with other elements of nature (Mattalia et al., 2025, in review). We follow the designation of Mattalia et al., (2025) in having three subcategories. Stewardship of target species (2.2.1) can be seen in the Solomon Islands, such as practices by Uluna and Sutahuri tribes in Guadalcanal. Members of these tribes enact stewardship of the eel mauvo or puleo ( Anguilla marmorata ), refraining from harvesting them and strictly protecting them within their waterways (Hevalao unpub. data). Similarly, this species is an aquatic animal totem for Delailasekau village of the Naitasiri highlands of Fiji and therefore there is a taboo for killing and eating them across Fiji Islands (Rashni & Mailautoka, unpub. data). The Menominee Nation, USA, has a long history of stewardship of the culturally important nämä'o or lake sturgeon ( Acipenser fulvescens ) (Fig. 2 ), which is evident today in recovery of the species in the Winnebago System as a result of co-management by the Menominee and others 78 . The Yakel tribes of Tana islands, Vanuatu, maintain strong Kastom beliefs and protect sacred sites, including freshwater springs and river sources, demonstrating stewardship of an assemblage of species (2.2.2) such as Anguilla spp. (eels) and Sicyopterus spp. (gobies). Water sources are often tied to ancestral spirits, with certain areas tabu (off-limits) for extraction or development of waterways (Hevalao unpub. data), which is stewardship of water and land (2.2.3). This can also be illustrated by the Nimiipuu People, whose water stewardship practices (affecting salmon) include knowledge of watershed protection and ongoing advocacy for breaching of the four lower Snake River dams that currently inhibit the free-flowing rivers required by native salmonids 79 . Domain 3: Knowledge Systems This domain concerns the structure of knowledge systems and knowledge transmission across and within generations (Fig. 1 ). While all categories in this framework represent types of knowledge (from knowledge of recipes to religion), this domain specifically focuses on the structure, storage, and transmission of knowledge. This may involve knowledge about the species themselves, or instances where the species play a role in transmitting other kinds of knowledge. Fundamental to the holding and transmission of species-related knowledge is its naming (3.1, names ): folk names and taxonomies are a culturally-rooted system of classification, conveying knowledge about the species and its cultural conception 47 , 80 . For example, Anguilla anguilla is ‘eel’ in the English language and English folk taxonomies (3.1.1). The immature life stage during which eels move upriver from the sea is called ‘elver’, a corruption of the earlier ‘eelfare’ meaning ‘passage of young eels upriver’. This name is part of the vocabulary of nature (3.1.2) as it transmits (or transmitted) knowledge about the species, as is the Japanese name, ito, for the Sakhalin taimen (Para hucho perryi) , indicated in Kanji as 伊富. It is written using character parts for "fish" and "ogre/demon", reflecting its folkloric role as a monster in the water. The category alternative word usage (3.1.3) includes instances when a taxon’s name also has other uses, such as ‘catfish’ in modern English: both the folk name of the Siluriformes and a term for online identity deception 81 . The category observations (3.2) includes observations of species behaviour, life history, or ecology that are transmitted or interpreted culturally, such as through traditional or Indigenous knowledge. Some South African Indigenous Peoples consider bullfrogs ( Pyxicephalus spp.) to rain from the sky, since these frogs spend most of the year aestivating underground, unseen until they emerge during heavy rains at the height of the rainy season 19 . Species can also be important in knowledge acquisition (3.3) such the African clawed frog ( Xenopus laevis ), an important model organism for biology education 82 , thereby playing a key role in knowledge transmission (3.4). Knowledge transmission can take many forms, such as in the São Benedito community, Ilha do Ituqui, Brazil, where fishing for pirarucu ( Arapaima gigas ) is associated with various knowledges, and the practice of pirarucu fishing enables personal and social transfer or development of these knowledges 83 , 84 . A similar role is conferred through the importance of salmon ( Oncorhynchus spp.) to the Nimiipuu People, for whom “the annual return of the salmon allows the transfer of traditional values from generation to generation” (Columbia River Inter-Tribal Fish Commission, 2006, p. 4) 79 . Species can play a role in knowledge inspiration (3.5), such as the electric eel ( Electrophorus electricus ), a key inspiration for the development of batteries by the European scientific community 85 . They learnt of this knowledge from observations (3.2) from local fishers near Calabozo, modern day Venezuela, likely Indigenous Peoples. Domain 4: Worldviews, Belief Systems, and Cultural Identities This domain includes human-freshwater fauna interactions that reflect a species’ role in worldviews, belief systems, and identity formation (Fig. 1 ). This includes species inclusion in socially constructed frameworks, such as religious or spiritual beliefs, and the oral or written traditions that support them, including myths, legends, and histories. This domain also includes the contribution of species to cultural or regional identity. These roles are often deeply embedded in how cultures make meaning, relate to the environment, and maintain a sense of belonging. For example, Bäru, the saltwater crocodile ( Crocodylus porosus ), is considered protector of the Gumatj clan of the Yolŋu People of Arnhem Land, Australia 86 , a part of the Yolŋu worldview and belief system (4.1). Also in Australia, pondi, the Murray cod ( Maccullochella peelii ), is an important component of the Ngarrindjeri worldview (4.1), including the Ngarrindjeri philosophy of being (known as Ruwel Ruwar) 87 . Pondi is essential in the origin of the Murray River and in the Ngarrindje narrative account (4.3) of the Creation (Kaldowinyeri). A worldview and belief system (4.1) of the Nguni People in South Africa is that frogs, specifically their croaking, is a sign that the ancestors approve of a ritual that was performed at the river 88 , and in Fiji, when a totem species appears out of the ordinary in freshwaters, it is seen as a good omen to the tribal landowners (Rashni & Mailautoka, unpub. data). Many species have a role in spirituality and religion (4.2), such as taimen ( Hucho taimen ), believed in Mongolia to be children of the river god, linking people and the spirit of the land and water. The role of species in worldviews, religion and narrative accounts is not always positive. The Chinese Giant Salamander has a negative cultural value in some Chinese communities, associated with bad luck and especially dead children (salamanders being transformed dead babies, or eating them), leading to its common name ‘baby fish’ or 娃娃鱼 89 . For many communities in Amazonia, the river dolphin boto ( Inia spp.) is strongly associated with enchantment and shapeshifting (Fig. 2 ), often blamed for illegitimate pregnancies, the ‘children of the boto’. While the associated narrative accounts (4.3) are oftentimes taken light-heartedly, they have also been used to mask violence against women 90 . This domain captures aspects of cultural or regional identity and relationships (category 4.4). For example, lafaek, the saltwater crocodile features in the Timorese belief system, lulik, in Timor-Leste, where there is kinship (4.4.1) between certain cultural groups and the ‘ancestor crocodile’ or Avo Lafaek, the grandfather crocodile 91 . Koena, the Nile crocodile, is the totem of the Bakoena clans of Lesotho and South Africa 92 , 93 , therefore playing a role in supporting cultural identities (4.4.2). This even for the Bakoena who have moved to Lesotho, where crocodiles are not found. The crocodile is further a symbol of Lesotho’s national identity, as it features in the country’s coat of arms. Such relationships also provide terms for landscape, sense of place, and connection to place (4.4.3): kwena is the name for Crocodylus niloticus for the BaPedi People and is the place name for Mogalakwena municipality, and the Mogalakwena river, in South Africa’s Limpopo Province. Mogalakwena is a compound word referring to the might of the crocodile 94 . The domain also encompasses symbolic or associative meaning (4.5). Zulu people in South Africa use the general term for frogs as an idiomatic expression ( figurative language , 4.5.2) for nudity 95 , threats of violence, and being honest 96 . The crocodile or kwena to the Bafokeng clan of the BaTswana people, also has symbolic or associative meaning (4.5.1) as a symbol of peace, in addition to being their totem. The phrase “let the crocodile return to water” is used as a declaration of peace 97 (Fig. 2 ), an idiom, proverb, or turn of phrase (4.5.2). In Japan, the catfish, namazu 鯰 (primarily Silurus asotus ), is symbolic of earthquakes (4.5.1) and is used popularly and widely e.g., signs for roads closed after earthquakes. This can be explained by the worldview (4.1), popularly known although no longer literally believed that earthquakes are caused by the thrashing of a giant catfish living under the islands of Japan 98 . Species names can also be used to evoke meaning through associative naming (4.5.3). For example, the term ‘buaya darat’, meaning ‘land crocodile’, is used to describe men who are playboys in Indonesia or Malaysia, evoking the predatory nature of crocodiles. Domain 5: Cultural and Social Practices This domain contains categories that highlight how species are agents in shaping and maintaining cultures. It covers the involvement of fauna in social practices, ranging from everyday activities to celebratory rituals, social organisation, including gender roles and social stratification, and the rules or norms that influence relationships (such as laws, taboos, and resource use regulations). For example, the Amur carp ( Cyprinus rubrofuscus ), symbolic of determination and overcoming obstacles in Japan (4.4.4), is a key symbol in children’s day (previously boy’s day) in Japan. It is depicted in koinobori (carp streamers) (Fig. 2 ), part of category ceremonies, rituals, celebrations, and social practices (5.1). In the São Benedito community, Brazil, hunting of the pirarucu ( Arapaima gigas ) successfully gives fishers prestige in their communities 84 , an example of the relationship with a species contributing to social organisation (5.2). Fisheries, such as that in Tono Reservoir in Ghana, often lead to gendered roles in fisher communities 99 , another example of social organisation (5.2). Such gendered interactions with harvested freshwater species can have profound social impacts, including negative impacts, such as with Nile perch ( Lates niloticus ) in Lake Victoria communities. The migration of Nile perch leads fishers (typically men) to follow the species, increasing the likelihood of engaging in high-risk sexual activities while away from their spouses. At the same time, declining catches in the lake—particularly of native species—have led some fishmongers (typically women) to form sexual relationships with fishers to secure fish. Together, these factors have contributed to increased human immunodeficiency virus prevalence in fishing communities 100 . This domain also encompasses rules or customary norms (5.3), such as the well-known and popularly referenced law in the United Kingdom that the monarch owns all unmarked mute swans in open water in the country (Exchequer of Pleas 1592), or the rule that killing ngweṋam, the Nile crocodile, is taboo for the Vhavenḓa people 101 . Domain 6: Recreation, Leisure and Tourism The recreation, leisure and tourism domain encompasses interactions with a species – non-consumptive, consumptive, and also representational (i.e., when the species itself is not necessarily physically involved) – that are for the purposes of recreation, leisure, or tourism. For example, the Hecht or pike, Esox lucius , is the focus of recreational fishing (6.1) in Rügen, Germany, which includes both the consumption of the species and the capture of large ‘trophy’ individuals 102 . Its local cultural role is reflected, for example, in a playground where the see-saw is shaped and painted as a pike, a representation of the species in sports and games (6.2.2). Such representation can even be purely conceptual, such as the idea of a crocodile ( Crocodylus spp.) in the children’s game ‘Please Mr Crocodile’ played in various English-speaking cultures. This role of the species in traditional sports and games (6.2) can also involve the live animal (6.2.1), such as the fighting of siamese fighting fish ปลากัด, ( Betta splendens) in Thailand 103 (Fig. 2 ). Recreational practices can also include tourism (6.4.), such as cultural tourism (6.4.1), to visit the Chabdi barahi Temple and resident eels, in Nepal, or nature-based tourism (6.4.2) in Peru, where giant otter ( Pteronura brasiliensis ) sighting is a focus 104 . Species can also contribute to aesthetics (6.3), such as dragonflies ( Aeshnidae and Libellulidae spp.) being appreciated for their appealing aesthetics in urban London, UK 105 . Domain 7: Creative Expression The final framework domain captures the expression of representations of the species in various creative aspects: visual, performing, and language arts. Such expressions are very often a reflection of the significance of species in the other six domains, as artists choose to use the arts to express different roles of species in their culture. For example, the saltwater and freshwater crocodiles of Australia ( Crocodylus porosus and C. johnstoni ), bäru and ngäw, are figures in the artworks (7.1.1) of the Yolŋu people of Australia, with stylistic choices differentiating these culturally important species 86 . Knowledge of the two species is also conveyed through songs ( music , 7.2.1.) and dance (7.2.2), where bäru is depicted with arms extended in front of the body and ngäw by arms tucked in beside the body 86 . Folksongs of South Africa’s Venda and Tsonga cultures include frogs in lyrics 106 , 107 . This is taken further in Pedi culture by mimicking the frogs croak, while singing those folks songs. Visual arts (7.1) include representations of the species in multiple respects, for example in crafts, artefacts, and tools (7.1.3), such as a West African Akan goldweight shaped like the eel, Anguilla mossambica (Fig. 3 ) (Sheales 2014). In Betawi communities in Indonesia, the saltwater crocodile is represented in culinary preparations (7.1.5) of Roti Buaya ‘bread crocodile’. The bread is symbolic of the commitment of the groom to act like the crocodile, who is believed by Betawai people to be faithful to its partner, and protect their children and partner from harm 108 . In Finland, thousands of years of cultural importance of the hauki or pike ( Esox lucius )(Fig. 2 ) can be seen through: prehistoric cave artworks (7.1.1.) of the species acting as spirit guide to a shaman 109 ; through important roles of the species in the Finnish national Epic Kalevala, a compilation of epic poetry (7.3.2) and one of the most significant works of Finnish literature (7.3.1); through to modern popularity in recreational fishing and as food, reflected in artwork ‘The pike is a fish’ by artist Outi Piiroinen in her series ‘Ode to Finnish food fish’ 110 . Overlap in domains, and differing or conflicting relationships Many of the species used as examples are culturally significant in specific cultures, with relevance spanning multiple domains. For example, the consumption of a species in ritual or social practices ( consumptive use , 1.1.4) evidently overlaps with social practices, rituals, and festive events ( cultural and social practices , 5.1), such as eating the Ilish fish on the first day of the Bengali year (1.1.4 and 5.1) 57 . Other categories and domains also overlap in some cases. For example, the use of the Siamese fighting fish in Thailand (Fig. 2 ), part of the domain recreation, leisure and tourism , has led to the species being popular in companionship (1.6) as well as in supporting cultural identities (4.4.2 in worldviews, beliefs and identities ) as the National Aquatic Animal of Thailand 103 . The species was inspiration for the national costume of Miss Universe Thailand in 2020, a role captured in the domain creative expression subcategories representation in costume, clothing, and jewellery (7.1.4) and other performing arts (7.2.5). Within some cultures, a given animal is immensely important across multiple domains, categories, and subcategories. This can be illustrated by considering some of the roles of Pacific salmon, chep, Oncorhynchus spp. in the culture of the Ainu of Hokkaido, Japan. The first salmon of the season is received in a ceremony known as Asircepnomi or Kamuycepnomi (1.1.3) 111 . This is a ceremonial ritual (5.1), which falls in the domain of cultural and social practices , which also includes the customary rule (5.3) that salmon should be killed with a specific ceremonial stick ‘isapa-kik-ni’ 112 . This knowledge on how to correctly kill salmon, according to Ainu culture, also falls within the consumptive use category knowledge related to capture (1.2), specifically killing knowledge (1.2.3). This knowledge is conveyed between generations using songs 113 , a form of knowledge transmission (3.4) within the domain knowledge systems and transfer , but also music (7.2.1) a category within the creative expression domain. The songs convey a narrative account (4.3), which falls within the domain of worldviews, belief systems, and identities . In the narrative, Ainu people kill fish (Pacific salmon) with rotten sticks and, so, the fish god no longer sent the fishes to the Ainu: this would historically have jeopardised food security (1.1.1). The first salmon ceremony is now practiced again despite many decades of criminalisation, and there is ongoing activism to restore Ainu rights to traditional salmon fishing beyond these ceremonies 111 . The role of salmon in contemporary movements for Ainu rights and cultural resurgence highlights the importance of salmon fishing as part of Ainu cultural identity (4.4.2.). However, the salmon also have symbolic meaning (4.4.4) through their connection to Indigenous People’s rights 114 . There are many additional facets of salmon significance, such as the use of salmon skins in shoes (1.4.2) 115 . Some species have different roles in different communities, which can lead to conflict, particularly when consumption (especially overconsumption) of a species might jeopardise roles in other communities. For example, the diamondback terrapin ( Malaclemys terrapin ) is Maryland’s state reptile and mascot for the University of Maryland ( supporting cultural identities 4.4.2.). However, this role was threatened when the species was heavily consumed as a delicacy (1.1.3.) up until the 1990s, leading to its overexploitation and heightened risk of extinction; despite prohibition, the species was still being harvested as recently as 2016 116,117 . Such conflicting interactions can lead to profound cultural losses, such as the historical extirpation of the namä'o or lake sturgeon in the Menominee Indian Reservation. Namä’o has a key role in cultural identity (4.4.2) including in the Metawin religion of the Menominee ( spiritual or religious beliefs 4.2), in the creation story ( narrative accounts 4.3), as a clan totem ( kinship 4.4.1), as protector of rice beds, and as the tribal historian, keeping stories throughout its long life ( worldviews and belief systems 4.1) 118 , 119 . Throughout history the nämä’o was important for Menominee food and nutritional security (1.1.1,1.1.2), and traditional medicines (1.3.1). The sturgeon was called by the sound of beating drums (an observation 3.2 of the sound ‘sturgeon thunder’ hundreds of years before Western scientific documentation of the sound 120 ), to Namä'o Uskíwämît (meaning sturgeon spawning place, vocabulary of nature 3.1.2.). The first sturgeon is celebrated with a welcoming and feast in celebration and thanks for the arrival of the sturgeon ( consumption in ritual and social practices 1.1.4, social practices, ritual and festive events 5.1) 118 , 119 . However, commercial fishers from US settler communities historically viewed the lake sturgeon as a nuisance species, and therefore many sturgeons were persecuted through non-use removal (1.8) or possibly for use in steamship boilers ( energy , 1.5.1) 121 . Latterly, settler communities extensively overconsumed the species when it became valuable for its smoked meat and caviar ( delicacies 1.1.3) and isinglass ( food processing and preservation 1.1.6 and material in crafts, artefacts or tools 1.4.1) 121 . Overexploitation and dam building led to collapses in sturgeon populations, and the extirpation of the nämä'o in the Menominee Indian Reservation. This was a tragedy for the Menominee nation who had already had most of their land taken by settlers, and had been limited to the modern site (2.1.3) of the Menominee Indian Reservation through a treaty in 1854. A major reason this land was chosen by the Menominee was it contained Namä'o Uskíwämît 118 . This loss was described as “very cultural and spiritual loss, loss of food sustenance, loss of cultural identity” (David Nahwahquaw Grignon 119 ). The importance of the nämä'o was not forgotten, and stewardship by the Menominee Nation ( stewardship of target species 2.2.1), including in collaboration with other groups such as recreational fishers (6.1), has led to the restoration of sturgeon populations in the system, and the return of the species to Namä'o Uskíwämît 78 . Discussion Our novel framework provides a basis for systematically and hierarchically organising and analysing human-freshwater fauna interactions, structured into seven cohesive yet interconnected domains. It has the potential to serve applications in conservation policy, implementation and communication, sustainable management, interdisciplinary research, and the preservation of natural and cultural heritage. We provide a comprehensive lens for understanding a spectrum of human- fauna interactions that can be used to broaden utilitarian approaches to understanding and managing the natural world. This lens can help incorporate the relational values, social meanings, and intangible connections that are all too often missing from the discourse around freshwater biodiversity and ecosystem management. Insights into human-freshwater fauna relationships Our results show that, across cultures, there are multiple, complex and sometimes conflicting interactions between people and species of freshwater fauna, informing a broad range of very different cultural relationships in different communities. Such interactions range from consumptive uses to the role of species in framing worldviews. In this framework we moved away from binaries such as ‘material and immaterial’ and ‘tangible and intangible’, as such dualisms can artificially segregate meaning and practice. We argue that the use of our framework provides room for a more comprehensive understanding between people and freshwater fauna. For example, the relationships between Sindhi people and the fish palla could be collapsed down to ‘provision of food’ when using the ecosystem services framework. In contrast, our framework offers a method to resist such oversimplification: palla are important in foodways, and are sacred, they are pilgrims who must be able to undertake their pilgrimage (which is also their migration), they are favoured as an offering to the saint Khwaja Khizar 122 , and are served as a sign of honour and respect to guests. The framework also offers a way to see how different versions of sacredness and respect for species are shown in different cultures: to some, a sacred species is one that should not be killed, such as the crocodile Bäru for the Yolŋu people in Australia 86 , to others, a sacred species is one that is used, but with respect and restraint for the gift of their lives, such as the lake sturgeon, namä’o for the Menominee people in the USA 118 . Different worldviews shape different relationships with other-than-human nature, and each relationship can only be fully understood within its specific cultural context. However, our framework provides a valuable tool for recognising these interactions from both external and global perspectives. The framework supports cross-cultural communication and understanding of human-fauna interactions in freshwater ecosystems, whether they occur in societies different from, or inclusive of, our own. Furthermore, the examples highlight how culture significance is sometimes tied directly to a single scientific species, and other times a broader range of taxa are referred to by a given ethnospecies name, and therefore cultural significance and the scientific species concept are not always congruent. Our examples and framework illustrate that the impacts of interactions of people and freshwater species can be positive, negative or neutral for people and/or other components of nature. These positive and negative contributions can be seen as parts of nature’s contributions to people, one of the frameworks with which we have aligned our work 25 . For example, the relationship between Finnish people and pike, hauki, is one in which pike provides positive contributions to people: from foodways to recreational fishing, to roles in narratives including folktales and the national epic, a feature in art from prehistoric caves paintings to modern design, interactions with pike shape and enrich Finnish culture (Fig. 2 ). Conversely, some relationships result in negative contributions to people, for example the changes to the ecosystem and associated fisheries in Lake Victoria after Nile perch introduction, with the consumptive relationship with the species in combination with its role in social organisation lead to profound negative social and public health impacts 100 . Particular interactions may have different impacts on different people in a community, such as the beliefs and associated narratives of the boto encantado (supernatural river dolphins) in Amazonia being valued stories for exploring concepts of cultural change and identity, but also being used to mask sexual violence 90 . Our results also highlight influence in the other direction, of ‘people’s contributions to nature’. Positive contributions of people to nature are perhaps most clearly seen in the category of stewardship practices (2.2) which follows Mattalia et al. , (2025, in review), but closely aligns with cultural values of species that are captured in other domains. For example, the positive impact of the Uluna and Sutahuri tribes on the eel mauvo or puleo, who see eels as guardians of river health, and in turn enact stewardship practices such as strictly protecting them, or the Nimiipuu people whose activism to push for breaching dams on the lower Snake river contributes to the stewardship of migratory salmonids 79 . Such examples of positive contributions of people to nature are abundant among Indigenous Peoples, but are not limited to them. An example from recreational fishers in Lower Saxony, Germany, is a transdisciplinary and co-developed project which investigated how different stewardship practices (targeted stocking vs whole ecosystem restoration) could enhance populations of fishes important to angling clubs and their members 123 . Negative contributions of people to nature can arise from interactions that benefit humans at nature’s expense—for example, the overconsumption of Chinese giant salamanders, has driven the species to near-extinction in the wild 60 . They can also stem from perceived negative interactions, such as non-use removal, which has led to local extirpations of alligator gar by U.S. recreational fishers who see the species as a competitor for prized game fish 72 . In sum, the framework we present and its illustrative examples highlight that the relationships between people and freshwater fauna are multidirectional, with contributions of people to other species, and from other species to people. The framework defines interactions and relationships, rather than uses and contributions, and in so doing considers this multidirectionality. This aligns with more recent broadening of the NCP approach to not just provide for contributions of nature to people, but also ‘bidirectional relationships that include reciprocity’ 124 , although the semantic lack of recognition of this in the term ‘nature’s contributions to people’ has been criticised 125 . The concept of reciprocity in our relationships with nature, including with water bodies and the (other) lives they contain, is central to many Indigenous worldviews, and also increasingly recognised by others as central to sustainability transformations 29 , 126 – 128 . Contribution of the framework to systematically organising and understanding different human freshwater-fauna interactions The seven domains are broad thematic groupings that facilitate our understanding of complex relationships between humans and freshwater fauna, and the role of ethnospecies in shaping the connection between culture, interactions and the concept of scientific species. The domain groupings were emergent from the more specific, and individually defined, categories. In some cases, interactions could be distinguished at a finer, subcategory scale. For example, distinguishing between fishing for food security (1.1.1), delicacies (1.1.3), or ritual consumption (1.1.4), in the category food and culinary heritage (1.1), allows capturing differing roles in culinary heritage, spiritual practices, or as a staple food, a delicacy, or a culturally significant ingredient. For some categories, such as narrative accounts, we found that a finer granularity was not possible for a universal framework. This is because the categorization of stories – such as whether they are seen to be histories, folktales, myths, or guidance – varies between cultures. A culturally specific application of the framework could, however, develop culturally appropriate subcategories. Our categories and subcategories are necessarily overlapping. Some subcategories always overlap with another category, such as examples placed in consumption in ritual and social practices (1.1.4) always also falling in social practices, rituals and festival events (5.1). Similarly, interactions may fall in multiple domains and categories for a particular human-fauna relationship. We highlight this inevitable overlapping as important in recognising that relationships with fauna can be plural, multifaceted, and difficult to compartmentalize. The fighting of pla kat or Siamese fighting fish ( Betta splendens ) is not simply a pet (consumptive use) or part of a game (recreation), but an important traditional cultural and social practice that is part of Thai national identity (worldviews and identity). Our finding that the relationship between a culture and species often includes many facets that fit into multiple distinct categories shows the plurality and complexity of considering the cultural significance of freshwater fauna. These examples highlight how many species have multiple and intersecting or amplifying importances to certain cultures. It is increasingly recognised that solutions to the global biodiversity crisis, most acutely exemplified in freshwater ecosystems, will require approaches that are transdisciplinary, weave multiple knowledges, accommodate diverse worldviews, and are Indigenous-led when in lands with Traditional Owners 22 , 126 , 129 , 130 . The framework presented here contributes to transdisciplinary approaches by connecting species (the unit typically used by natural scientists), to ethnospecies (the unit by which people conceptualise species), and to cultural significances in a systematic way that is conceptually simple to understand. The thematic and hierarchical structuring of the framework allows both an in-depth exploration of the multiple facets of the role of a given taxa in a particular culture as well as of the breadth of different types of importance between cultures, species, or globally. The multifaceted relationship between humans and eels ( Anguilla spp.) encompassing food security, religious and ritual practice, and cultural identity across the world, demonstrates how a single taxon can fulfil multiple cultural roles and situational meanings, and traverse political borders (Fig. 3 ). Additionally, the framework offers a way to approach documenting and understanding differing roles in different cultures, which may be contradictory or conflicting. For example, the overexploitation of lake sturgeon / namä’o by US settlers due to non-use removal and later view of them as food delicacies, combined with a lack of stewardship in settler culture, conflicted with the highly important role of namä’o in Menominee culture. This led to both negative impacts on the species and on a culture. Identification of contradictory relationships with the same species – or species in the same ecosystem – to different communities, is vital for understanding the conflicts that underscore much of biodiversity conservation and management of the natural world 131 . Of course, understanding of different interactions between people and fauna is just one part of conflict resolution, and occurs in the complex setting of trust, justice and power between the different communities involved 132 . This framework seeks to describe the interaction between people and freshwater animals, but we recognise that there are many values, importances, power imbalances, histories and human-human relationships that shape these interactions. Applications of the framework in interdisciplinary understanding The differences in language and terminology used by different disciplines can make interdisciplinary work challenging. The framework helps provide a ‘translating lens’ through which different types of scholarship can be understood more clearly, by providing a consistent framing vocabulary to the complexity of explaining culture. In the environmental humanities, for instance, approaches are often specified and relevant terminology used that natural scientists are unlikely to have come across, such as ‘posthumanism’, or ‘more-than-human’, but such work may contain in-depth discussions of human-species interactions relevant to a manager working with certain species. Similarly, fisheries scientists may write papers densely laden with terms such as ‘catch per unit effort’ or ‘stock recruitment relationship’ that may be alien to an environmental humanist. Both works may relate to the important role of a fish species in foodways and cultural identity, and therefore, the responsibility of communities to steward the species. This role could be outlined in a simple and clear way by applying our framework to these interactions, with reference to the more specific, disciplinary literature. Additionally, by categorizing significance from different cultures together, the framework can aid in cross-cultural understanding, as users can identify examples from their own culture, and use this to better recognize importance in other cultures. For example, the European eel is a culturally important food in the Netherlands (Fig. 3 ), but also a critically endangered species. For a Dutch person, understanding the cultural context of eating a critically endangered species (e.g., personal experiences of eating smoked eels with family) may help in building understanding and empathy as to why critically endangered species are also eaten in other cultures. Improving communication and understanding can also extend beyond preservation of biocultural diversity, into other areas, such as public health and medicine, which are also deeply embedded in cultural context. For example, small-scale gold mining leads to mercury contamination of water and then biomagnifies through trophic webs, creating public health and conservation concerns 133 , 134 . Dietary recommendations on which fish species are safe to consume – or not – through examining the nutritional, ecological, and market attributes, and mercury risks is a potential risk mitigation strategy 135 . A recent study shows that large, migratory fish in the Amazon (e.g. Brachyplatystoma rousseauxii ) have low nutrient density and high mercury content, and are also more threatened, and it suggested recommendations and messaging could discourage their consumption 135 . Understanding the cultural context, values and traditions around fish consumption, and integrating them with other information sources, could strengthen such an approach. In French Guiana certain species, including fish such as B. rousseauxii, Hoplias aimara , and crocodylians such as Paleosuchus trigonatus , have high mercury risks if consumed, but have a high cultural value for consumption (1.2.3), recreational hunting and fishing (6.1) and knowledge sharing of fishing techniques between generations (3.4), which can lead to continued consumption even in the knowledge of mercury risks. Application and policy implications of the framework in biodiversity conservation The framework proposed can complement ecology-oriented approaches for freshwater biodiversity conservation through different channels. Our biocultural approach highlights the bonds between freshwater fauna and various cultural groups in different regions and provides justifications for the protection of threatened freshwater fauna, beyond their ecological roles. Moreover, the biocultural framework can help gain momentum among the public for freshwater ecosystems to be experienced as vital parts of the environment upon which our society depends 136 . Freshwater ecosystems have often been regarded simply as a resource for clean water and food, with freshwater fauna such as fish, in many cases, not being considered wildlife, which could have negative impacts on the conservation of freshwater fauna 137 , 138 . By recognizing the cultural significance of freshwater fauna, researchers and conservation practitioners can work together to raise public awareness and increase their willingness to contribute to freshwater conservation through targeted communications that enhance emotional connections between human and freshwater fauna. Emotions, values and personal experience have been shown to be crucial for decision making, with emotionally driven storytelling demonstrably more effective than ‘fact-based’ narratives in shifting attitudes 139 . As conservation research and practice remain insufficient in the face of current threats to freshwater biodiversity, freshwater conservationists widely acknowledge that losses can only be stemmed by a paradigmatic shift in relationships between people and freshwater biodiversity 136 , 140 . Given the widely observed shifting baseline syndrome for freshwater fauna, where each generation accepts a more degraded ecological state as the baseline 141 , 142 , highlighting past human-freshwater fauna links and revitalizing lost cultural traditions and stewardship practices holds the potential for engaging younger generations in freshwater conservation and supporting more ambitious goals for restoration. In some instances, traditions may have long been obscured, for example the once important and multifaceted role of the eel in British society 143 . In such cases, an approach to understanding historical or obscure significance using the framework we outline can identify emotive and engaging local or national examples, to garner interest and engagement in freshwater conservation. In other instances, this may be a matter of preserving and supporting the revitalization of cultural traditions and practices that are being lost due to colonisation, cultural suppression, or social and environmental change. For example, in parts of Guadalcanal, Solomon Islands, the Uluna and Sutahuri tribes revere freshwater eels as guardians of river health, enacting stewardship practices such as refraining from harvesting them and strictly protecting them within their rivers and streams. Unlike in other regions where eels are a food source, this tribe views them as essential for maintaining river health and cleanliness. The eel’s role in the ecosystem aligns with their spiritual beliefs, reinforcing non-extractive conservation practices. Similarly, in Vanuatu, traditional fishing taboos regulate when and how certain species can be caught, ensuring sustainable use. By revitalizing these stewardship practices, younger generations can develop a stronger connection to freshwater biodiversity, reinforcing conservation efforts in both customary and scientific contexts. Such actions can be part of a broader place-based biocultural approach. For example, in the Solomon Islands, place-based conservation efforts—such as Locally Managed Marine Areas (LMMAs), and the iTina and Tina Rivers study which includes local rangers for boundary monitoring—integrate traditional ecological knowledge with modern conservation techniques, ensuring that freshwater resources are protected through a mix of Indigenous and scientific strategies, aligning conservation with cultural narratives and ancestral responsibilities. Future directions and next steps This framework to categorise the cultural significance of freshwater fauna provides an approach to conceptualise and organise interactions between people and freshwater fauna, and consequently perhaps the most direct application is to use the framework as a tool for collecting and understanding data on these interactions. For example, the framework could be used to develop more comprehensive and standardised assessments of the implications of freshwater biodiversity loss, or potentials of restoration, or it could be incorporated into tools to measure conservation impact, such as the Open Standards for the Practice of Conservation of the Conservation Measures Partnership 144 . The framework could be used as a basis for knowledge or database collation on culturally informed interactions between people and freshwater species in particular basins or regions or for particular taxonomic groups. For example, migratory fishes have been identified as needing conservation for their ecological and socioeconomic value, with the concept of identifying ‘Global Swimways’ proposed to address this need 145146 . However, so far, no social criteria have been suggested for Swimways, as they are seen as difficult to quantify 146 . The framework presented here provides a systematic method to approach this issue. Similarly, freshwater megafauna have been identified as a group in particular need of urgent conservation action 147 , and as having strong potential to serve as freshwater flagship species 148 , a role that is especially important given repeated calls to identify such species to help raise the profile of their threatened aquatic ecosystems 136 , 140 . However, flagship species operate as cultural assets, particularly through their presence in pre-existing cultural framings, alongside alignment with current cultural trends and political conditions 149 . This framework could provide a more effective approach to identifying potential flagship species, as well as for identifying with which communities a species may, or may not, galvanise action. Our focus in this research has been on species of freshwater fauna, due to the particularly acute status of freshwater biodiversity, and the highly interrelated relationships between humans and all aspects of freshwaters and freshwater life. However, many of the species in our examples are migratory or multi-realm occupants, and spend part of their lives in marine systems. Given that relationships with such species do not stop at the coast, nor should consideration of their cultural roles in management or conservation. Investigations into how well the framework can be applied more broadly to all aquatic fauna, or to flora and funga, and beyond aquatic habitats to those on land, are interesting future avenues to be explored. Finally, to borrow from Kimmerer’s (2012, p. 319) 29 thinking on teaching traditional ecological knowledge as a “prompt to the imagination, a mirror in which western society can examine its own relationship to the living world”, perhaps our framework can contribute to introducing and inspiring people who do not currently have reciprocal relationships with freshwater fauna to reconsider their own relationships and interactions. By presenting other ways of relating, and providing a way to systematically consider these as a complement to scientific thinking does not try to separate the tangible and intangible, our work could be a prompt to imagine new ways of relating to, and of conserving, freshwater species, such that these interactions are possible into the future. Declarations Author Contributions PG, RET, FH, SK, PP and SCJ conceived the initial concept, which was initially developed by RET and PG with inputs from FH, SK, PP and SCJ. Example collection and contribution was conducted by PG, RET, FMP, LKFC, FH, RSH, SK, KKM, and BR. Framework definitions were developed by PG, RET and FMP, with extensive input from other authors. Framework refinement and completion was led by PG, with all authors contributing. PG took the lead in writing the manuscript, with all authors contributing sections, comments, or refinements to multiple drafts. All authors provided critical feedback and helped shape the research, analysis and manuscript. Acknowledgements This work has been supported by the Leibniz Competition project ‘Freshwater Megafauna Futures’ to SCJ and PG. RT recognises the research fellowship support of LE STUDIUM Loire Valley Institute for Advanced Studies, Orléans and Tours, France, with funding from the European Union’s Horizon 2020 research and innovation programme under Marie Skłodowska-Curie grant agreement No. 665790, for a project on ‘Biological and cultural diversity, and flow regimes – a unifying approach for managing riverscapes’ which helped catalyse the collaboration reflected in this paper. RET further acknowledges funding support from The Nature Conservancy in framework drafting. FH acknowledges the support from the Chinese Academy of Sciences (E355S122). SK acknowledges the funding support from an anonymous donor to The Nature Conservancy. VRG acknowledges support from the “María de Maeztu” Programme for Units of Excellence of the Spanish Ministry of Science and Innovation (CEX2019-000940-M). 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Lemaire, J. Using Crocodylians for monitoring mercury in the tropics. Ecotoxicol. 2023 328 32 , 977–993 (2023). Heilpern, S. A. et al. Accessible, low-mercury, and nutritious fishes provide win-wins for conservation and public health. One Earth 8 , (2025). He, F., Jähnig, S. C., Wetzig, A. & Langhans, S. D. More exposure opportunities for promoting freshwater conservation. Aquat. Conserv. Mar. Freshw. Ecosyst. 31 , 3626–3635 (2021). Wyatt, T., Friedman, K. & Hutchinson, A. Are Fish Wild? Liverp. Law Rev. 42 , 485–492 (2021). Tian, M., Potter, G. R. & Phelps, J. What is “wildlife”? Legal definitions that matter to conservation. Biol. Conserv. 287 , 110339 (2023). Toomey, A. H. Why facts don’t change minds: Insights from cognitive science for the improved communication of conservation research. Biol. Conserv. 278 , 109886 (2023). Kalinkat, G. et al. Flagship umbrella species needed for the conservation of overlooked aquatic biodiversity. Conserv. Biol. (2017) doi:10.1111/cobi.12813. Humphries, P. & Winemiller, K. Historical impacts on river fauna, shifting baselines, and challenges for restoration. Bioscience 59 , 673–684 (2009). Kochalski, S., Riepe, C., Fujitani, M., Aas, Ø. & Arlinghaus, R. Public perception of river fish biodiversity in four European countries. Conserv. Biol. 33 , 164–175 (2019). Greenlee, J. W. Seeing all the Anguilles: Eels in the cultural landscape of medieval and early modern England. (Cornell University, 2020). Redford, K. H., Hulvey, K. B., Williamson, M. A. & Schwartz, M. W. Assessment of the Conservation Measures Partnership’s effort to improve conservation outcomes through adaptive management. Conserv. Biol. 32 , 926–937 (2018). Stoffers, T., Sayer, C. A., Baratech, A. & He, F. ‘Global swimways’ on free-flowing rivers will protect key migratory fish species. Nature 627 , 489 (2024). Worthington, T. A. et al. Global Swimways for the conservation of migratory freshwater fishes. Front. Ecol. Environ. 20 , 573–580 (2022). He, F. et al. The global decline of freshwater megafauna. Glob. Chang. Biol. 25 , 3883–3892 (2019). Carrizo, S. F. et al. Freshwater Megafauna: Flagships for Freshwater Biodiversity under Threat. Bioscience 67 , 919–927 (2017). Jepson, P. & Barua, M. A Theory of Flagship Species Action. Conserv. Soc. 13 , 95–104 (2015). Additional Declarations There is NO Competing Interest. Supplementary Files SupplementalInformationTablesS1S2S3.pdf Supplementary information tables S1, S2 and S3 Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6830897","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":474177076,"identity":"7403a1f4-ebe6-44b7-994a-41be05fb1dcc","order_by":0,"name":"Phoebe Griffith","email":"data:image/png;base64,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","orcid":"https://orcid.org/0000-0001-7274-0137","institution":"Leibniz Institute of Freshwater Ecology and Inland Fisheries","correspondingAuthor":true,"prefix":"","firstName":"Phoebe","middleName":"","lastName":"Griffith","suffix":""},{"id":474177077,"identity":"ac857046-57de-40b0-9d95-04d2506c19e1","order_by":1,"name":"Sonja Jähnig","email":"","orcid":"https://orcid.org/0000-0002-6349-9561","institution":"Leibniz Institute of Freshwater Ecology and Inland Fisheries","correspondingAuthor":false,"prefix":"","firstName":"Sonja","middleName":"","lastName":"Jähnig","suffix":""},{"id":474177078,"identity":"61cc0911-5abb-4a56-a68c-fe8677bdcd9d","order_by":2,"name":"Rebecca Tharme","email":"","orcid":"","institution":"Riverfutures","correspondingAuthor":false,"prefix":"","firstName":"Rebecca","middleName":"","lastName":"Tharme","suffix":""},{"id":474177079,"identity":"429134b2-7f4c-448a-ab53-245f60971da2","order_by":3,"name":"Fortunate Phaka","email":"","orcid":"","institution":"African Amphibian Conservation Research Group","correspondingAuthor":false,"prefix":"","firstName":"Fortunate","middleName":"","lastName":"Phaka","suffix":""},{"id":474177080,"identity":"8e132fff-56d1-4b7c-a1bc-fc1156cc8d04","order_by":4,"name":"Lekima Copeland","email":"","orcid":"","institution":"The University of the South Pacific","correspondingAuthor":false,"prefix":"","firstName":"Lekima","middleName":"","lastName":"Copeland","suffix":""},{"id":474177081,"identity":"bfc1bffb-29ab-4f53-9628-3763c77ad0b7","order_by":5,"name":"Fengzhi He","email":"","orcid":"https://orcid.org/0000-0002-7594-8205","institution":"Northeast Institute of Geography and Agroecology, Chinese Academy of Sciences","correspondingAuthor":false,"prefix":"","firstName":"Fengzhi","middleName":"","lastName":"He","suffix":""},{"id":474177082,"identity":"3ea52452-6af4-4f6c-ab3e-a3e4a4218108","order_by":6,"name":"Robson Hevalao","email":"","orcid":"","institution":"Islands Resources Nexus","correspondingAuthor":false,"prefix":"","firstName":"Robson","middleName":"","lastName":"Hevalao","suffix":""},{"id":474177083,"identity":"789d8603-f3ac-4d24-8b8c-8010fa49b793","order_by":7,"name":"Shiteng Kang","email":"","orcid":"https://orcid.org/0000-0001-8326-0286","institution":"The Nature Conservancy","correspondingAuthor":false,"prefix":"","firstName":"Shiteng","middleName":"","lastName":"Kang","suffix":""},{"id":474177084,"identity":"96bc6758-4ef3-4e40-b121-612203e1460c","order_by":8,"name":"Simone Langhans","email":"","orcid":"","institution":"Otago Regional Council (ORC)","correspondingAuthor":false,"prefix":"","firstName":"Simone","middleName":"","lastName":"Langhans","suffix":""},{"id":474177085,"identity":"1703dd01-de5f-4176-b77e-5023b853b9ed","order_by":9,"name":"Kini Mailautoka","email":"","orcid":"","institution":"National Trust of Fiji","correspondingAuthor":false,"prefix":"","firstName":"Kini","middleName":"","lastName":"Mailautoka","suffix":""},{"id":474177086,"identity":"ffb91b69-b665-4482-b4e5-6ab32847f5a3","order_by":10,"name":"Paolo Petry","email":"","orcid":"","institution":"The Nature Conservancy","correspondingAuthor":false,"prefix":"","firstName":"Paolo","middleName":"","lastName":"Petry","suffix":""},{"id":474177087,"identity":"542e0999-7fd4-4c31-a82f-f06454d7b983","order_by":11,"name":"Bindiya Rashni","email":"","orcid":"","institution":"University of the South Pacific","correspondingAuthor":false,"prefix":"","firstName":"Bindiya","middleName":"","lastName":"Rashni","suffix":""},{"id":474177088,"identity":"11e4ee54-0c89-43bf-b9c1-56a1ef2509fd","order_by":12,"name":"Victoria Reyes-Garcia","email":"","orcid":"https://orcid.org/0000-0002-2914-8055","institution":"Institució Catalana de Recerca i Estudis Avançats","correspondingAuthor":false,"prefix":"","firstName":"Victoria","middleName":"","lastName":"Reyes-Garcia","suffix":""}],"badges":[],"createdAt":"2025-06-05 16:23:08","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6830897/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6830897/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":85212791,"identity":"0b966256-59ec-428e-92dc-4a87d6e9c807","added_by":"auto","created_at":"2025-06-23 12:43:45","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":498482,"visible":true,"origin":"","legend":"\u003cp\u003eVisualisation of the framework to categorise the cultural significance of freshwater fauna. The central doughnut displays the highest hierarchical level of the framework, the seven thematic domains. Outer circles, coloured by domain, show categories (inner ring) and subcategories (outer ring, where present). Some slight abbreviations of subcategory names have been made where necessary for visualisation.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-6830897/v1/6fbaa31ec65217e7a543b74d.png"},{"id":85211617,"identity":"4f992207-966a-420a-bd9c-9889306d8c61","added_by":"auto","created_at":"2025-06-23 12:35:45","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":560772,"visible":true,"origin":"","legend":"\u003cp\u003eIllustrations representing simplified aspects of different relationships between people and freshwater fauna. Clockwise, from top left: the palla is a prized food fish offered to honoured guests in Sindh, Pakistan; the phrase ‘let the crocodile return to the water’ is used as a declaration of peace by the Bafokeng Clan of the BaTswana People; the pike is popular in recreational fisheries in Finland, whilst also featuring in the national epic and in ancient cave paintings; worshippers at the pool of the Chabdhi Barahi temple in Tanahun, Nepal, offer sacrifices to the sacred eels as a conduit to, or incarnation of, the goddess; the sturgeon is of great cultural importance to the Menominee People, who have stewarded the species and instigated its return to historic spawning grounds; the Siamese fighting fish is popular in sports and aquaria in Thailand; the boto or river dolphin is an agent in narratives where it transforms into a man wearing white, and in a white hat, who seduces women; and streamers shaped like carp are flown on Children’s Day in Japan, as a sign of hard work and perseverance. Griffith (CC0)\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-6830897/v1/8312039b0d2188987861727a.png"},{"id":85213992,"identity":"6bea53e3-2208-4289-9a7d-74547c47f5a2","added_by":"auto","created_at":"2025-06-23 12:59:46","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":469253,"visible":true,"origin":"","legend":"\u003cp\u003eIllustrations of some of the culturally significant relationships between people and eel species (\u003cem\u003eAnguilla\u003c/em\u003e spp.) across the world. Eel traps are or have been woven to capture eels (\u003cem\u003e1.2.2\u003c/em\u003e) by the Nanticoke Indian Tribe, USA, as shown by the eel trap, and the British, as shown by the painting of eel pots on a bridge by artist William Collins (\u003cem\u003e7.1.1\u003c/em\u003e); the eel is featured in the coat of arms of Fockbeck, Germany (\u003cem\u003e4.4.2\u003c/em\u003e) as a key agent in a local folktale (\u003cem\u003e4.2\u003c/em\u003e); a work (\u003cem\u003e7.1.1\u003c/em\u003e) by Japanese artist Katsushika Hokusai illustrates the idiom ‘unagi nobori’ (eel climbing, to climb inexorably, \u003cem\u003e4.5.2\u003c/em\u003e) as fishers (\u003cem\u003e1.1.3)\u003c/em\u003e failing to keep their catch of giant eels; an eel spear (\u003cem\u003e1.1.1, 1.1.4, 1.2.2\u003c/em\u003e) from the Mi’kmaq Nation, Canada; a beaded eel (\u003cem\u003e7.1.1\u003c/em\u003e) by artist Lisa Thomas Boots, an image of the American eel as Clan Animal of the eel clan (\u003cem\u003e4.4\u003c/em\u003e) of the Onondaga Nation, USA; a 1994 stamp (\u003cem\u003e7.1.3\u003c/em\u003e) showing Māui fighting the eel god Tuna, part of the narrative (\u003cem\u003e4.3\u003c/em\u003e) explaining the origin of eels (\u003cem\u003e4.1\u003c/em\u003e) in Aotearoa/New Zealand; a restaurant mural (\u003cem\u003e7.1.1\u003c/em\u003e) of the goddess of the Chabdi Barahi temple Nepal (\u003cem\u003e4.2\u003c/em\u003e) with her protector/messenger the eel; an Akan goldweight representing an eel (\u003cem\u003e7.1.3\u003c/em\u003e), part of the Akan measuring system for fair-trade in Ghana; eels are being smoked to prepare them for being sold as a popular delicacy (1.1.3) in the Netherlands. Copyright of images, clockwise from far left: ©Victoria and Albert Museum, London; \u003cdel\u003e©\u003c/del\u003eGriffith; \u003cdel\u003e©\u003c/del\u003ePublic Domain; \u003cdel\u003e©\u003c/del\u003ePublic Domain; ©Canadian Museum of History; ©Smithsonian National Museum of the American Indian; ©Carnegie Museum of Natural History; \u003cdel\u003e©\u003c/del\u003ePublic Domain; \u003cdel\u003e©\u003c/del\u003eGriffith; ©The Trustees of the British Museum\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-6830897/v1/7555ec960ebcf8d01ae4f394.png"},{"id":100364166,"identity":"ef109153-a3c7-4ed9-aa0f-851058914f43","added_by":"auto","created_at":"2026-01-16 07:52:45","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2939395,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6830897/v1/741b8143-d5f8-48f7-8a20-e078fac3331c.pdf"},{"id":85211618,"identity":"e00b4545-50d7-426d-a488-991e4ca42ff4","added_by":"auto","created_at":"2025-06-23 12:35:45","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":1268125,"visible":true,"origin":"","legend":"Supplementary information tables S1, S2 and S3","description":"","filename":"SupplementalInformationTablesS1S2S3.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6830897/v1/2d499800837f4cdcfea0f07c.pdf"}],"financialInterests":"There is \u003cb\u003eNO\u003c/b\u003e Competing Interest.","formattedTitle":"A framework to categorise the cultural significance of freshwater fauna","fulltext":[{"header":"Introduction","content":"\u003cp\u003eInland waters have played an essential role in human societies since time immemorial, with freshwaters \u0026ndash; including rivers, lakes and wetlands \u0026ndash; shaping the history of humanity\u003csup\u003e\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e,\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e\u003c/sup\u003e. Even today, 90% of the world\u0026rsquo;s population lives within 10 km of surface freshwater bodies\u003csup\u003e\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e\u003c/sup\u003e. The disproportionately high biodiversity of freshwaters, relative to their small proportion of the earth's surface, is well established\u003csup\u003e\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e\u003c/sup\u003e. Freshwater ecosystems are also known for their formative and critical roles in the lifeways of peoples of different cultures\u003csup\u003e\u003cspan additionalcitationids=\"CR6\" citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e\u003c/sup\u003e. Consequently, freshwaters have high biocultural diversity and value. Biocultural diversity explicitly recognises the deeply intertwined, co-evolved and dynamic relationships between biological and cultural diversity, and the often place-based relationships that underpin socio-ecological systems\u003csup\u003e\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e,\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u003c/sup\u003e. Nonetheless, both biological and cultural diversity are imperilled. One quarter of freshwater fauna species are threatened with extinction\u003csup\u003e\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e\u003c/sup\u003e, with the potential impacts of this loss on the human cultures and communities they interact with extending far beyond a count of threatened species. Cultural diversity is also at risk, with nearly half of the world\u0026rsquo;s spoken languages on the brink of extinction\u003csup\u003e\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e\u003c/sup\u003e. The loss of these languages is closely tied to both cultural and biodiversity declines\u003csup\u003e\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e\u003c/sup\u003e. Simultaneously, erosion of relationships between people and freshwater biodiversity both exacerbates species extinction risks and negatively impacts human wellbeing\u003csup\u003e\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e,\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e\u003c/sup\u003e, particularly through the loss of Indigenous and other local knowledges\u003csup\u003e\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eAs much of the biodiversity that exists today has co-evolved with human cultural diversity, in many contexts their maintenance is inseparable\u003csup\u003e\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e,\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u003c/sup\u003e. For biodiversity conservation, management and governance to be ethical, equitable and effective, therefore, cultural context should be a central consideration\u003csup\u003e\u003cspan additionalcitationids=\"CR18\" citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e\u003c/sup\u003e. Such consideration is likely to be particularly key in freshwater environments, where humans and nature are deeply entwined\u003csup\u003e\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e\u003c/sup\u003e. However, the vital role of culture in conserving biodiversity \u0026ndash; and of biodiversity in sustaining cultures \u0026ndash; has yet to be fully acknowledged in the global discourse on environmental protection\u003csup\u003e\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e\u003c/sup\u003e. In fact, while the multi-directional relationships and nature-culture nexus are well conceptualised in some worldviews, particularly those of many Indigenous Peoples\u003csup\u003e\u003cspan additionalcitationids=\"CR23\" citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e\u003c/sup\u003e, the complex and culturally informed relationships between people and other elements of nature have been largely overlooked in the Western scientific worldview. Consequently, until recently, relations between nature and culture have been largely absent in international and national policy instruments addressing the biodiversity crisis\u003csup\u003e\u003cspan additionalcitationids=\"CR26\" citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e\u003c/sup\u003e. This situation is changing, however as reflected in the Convention on Biological Diversity Decision 15/22 on Nature and Culture\u003csup\u003e\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e\u003c/sup\u003e and the adoption of the concept of nature\u0026rsquo;s contributions to people (NCP) by the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES). The IPBES defines NCP as \u0026ldquo;all the contributions, both positive and negative, of living nature to people\u0026rsquo;s quality of life\u0026rdquo; (D\u0026iacute;az et al., 2018, p. 270), recognising the critical role of culture in defining all relationships between people and other nature. Such changes are important, as many sustainable solutions to environmental degradation are found at the intersection of nature and culture, in the form of reciprocal relationships with other-than-human nature\u003csup\u003e\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eConceptual and applied frameworks aid in understanding and describing the types of diverse relationships that exist between people and freshwater biodiversity. Such frameworks help communicate the context-specific cultural interactions between people and nature to those in other communities, aiding in global uptake and understanding of biocultural approaches\u003csup\u003e\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u003c/sup\u003e. For example, the Ramsar Convention, a critical policy instrument for freshwater ecosystems, has recognized the importance of culture since the treaty was adopted in 1971\u003csup\u003e6\u003c/sup\u003e, and has set out a typology of significant wetland-related activities to systematise the types of cultural significance wetlands, as places, hold for people\u003csup\u003e\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e\u003c/sup\u003e. Using a different lens, Lynch \u003cem\u003eet al.\u003c/em\u003e (2023) applied the IPBES NCP reporting categories to identify various contributions of freshwater biodiversity to people, including non-material cultural contributions. However, in their application, \u0026lsquo;culture\u0026rsquo; is treated as a singular undifferentiated category, separate from other culturally rich categories, such as food or recreation, as is also the approach in the Ecosystem Services framework\u003csup\u003e\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e\u003c/sup\u003e. Interpretation of the contribution of freshwater species thus seems to depend on the concept of culture as considered by those applying the framework, with potential for a narrow or biased scope. While these kinds of frameworks have progressed, we still lack a robust, unifying framework for categorising and systematically understanding, including for practical application, the importance that animal species found in freshwaters may hold for different communities and cultural groups.\u003c/p\u003e \u003cp\u003eThe absence of a such framework for assessing the cultural significance of freshwater species is particularly problematic in that species remain a primary unit for biodiversity conceptualisation and practical conservation action\u003csup\u003e\u003cspan additionalcitationids=\"CR32\" citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e\u003c/sup\u003e. Biocultural approaches that consider the place-based interrelationships of people and nature at the species level are useful for efforts in transdisciplinary conservation\u003csup\u003e\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u003c/sup\u003e. Consequently, a framework for connecting the scientific concepts of species (or other taxonomic groups or taxa) to their social and cultural contexts can be a pragmatic tool for knowledge sharing across disciplines and among cultures. Such a framework could be useful, for example, to communicate about migratory species that traverse multiple human communities, territories, and cultures. The conservation of such species and associated cultural diversity requires cooperation across these diverse groups. To illustrate, freshwater eels (\u003cem\u003eAnguilla\u003c/em\u003e species, spp.) are one such taxonomic group that migrates vast distances between fresh waters and oceanic breeding grounds to complete their life cycle\u003csup\u003e\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e\u003c/sup\u003e. The giant mottled eel (\u003cem\u003eA. marmorata\u003c/em\u003e) is crucial to communities across the Pacific, including in Melanesia, Polynesia and Kiribati\u003csup\u003e\u003cspan additionalcitationids=\"CR36 CR37\" citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e\u003c/sup\u003e. The species is an ecological indicator, and holds spiritual and economic significance\u003csup\u003e\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e,\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e\u003c/sup\u003e. In some areas, this species symbolizes ancestral connections, while in others, it is a totem species and or vital food source. A framework that allows regional communication and cooperation in managing these migrant populations would therefore be aided by mutual understanding of this diversity in human-eel relationships.\u003c/p\u003e \u003cp\u003eIn this context, we developed a first of its kind framework for capturing human-freshwater fauna interactions across diverse knowledge systems and disciplines, drawing on a broad and interdisciplinary literature. By categorising these relationships, we can conceptualise what cultural significance of species looks like practice. We define \u0026lsquo;cultural significance\u0026rsquo; as the importance of a particular taxon\u0026rsquo;s role within a specific culture\u003csup\u003e\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e,\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e\u003c/sup\u003e, including the values, importance, knowledge, meaning, and relationship of that species to a particular cultural group\u003csup\u003e\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e\u003c/sup\u003e. By proposing a framework to categorise different culturally informed interactions between people and freshwater fauna, we hope to provide a pragmatic way to improve understanding and strengthen the integration of cultural significance in safeguarding of natural and cultural diversity and heritage.\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eFramework Development and Category Definitions\u003c/h2\u003e \u003cp\u003eTo establish a unique framework to address the cultural significance of freshwater fauna, we first examined the ways in which elements of culture have already been framed and grouped, including for freshwater ecosystems. We developed the initial draft categories for our framework in large part through reviewing the categorisations used in five frameworks known to the authors: (i) the Ramsar Typology of Significant Wetland-Related Activities\u003csup\u003e\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e\u003c/sup\u003e, (ii) reporting categories for Nature\u0026rsquo;s Contributions to People\u003csup\u003e\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e\u003c/sup\u003e, (iii) UNESCO Intangible Cultural Heritage domains\u003csup\u003e\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e\u003c/sup\u003e, (iv) the International Union for Conservation of Nature (IUCN) Species Use Database\u003csup\u003e\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e\u003c/sup\u003e, and the (v) South African ethnoherpetological elements of culture\u003csup\u003e\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e\u003c/sup\u003e. We selected these frameworks either for their broad relevance to human interactions with freshwater species, and/or for their in-depth consideration of key aspects of cultural and use dimensions. We also chose them for their thematic diversity, established or emerging application in policy and practice, and/or finer-scale delineation of certain types of human-fauna relationships.\u003c/p\u003e \u003cp\u003eTo create an integrated and robust framework, categories and domains from each framework were compiled, reviewed for conceptual alignment, and consolidated or adapted, where duplicate or overlapping classifications existed. For example, some categories we directly drew from past frameworks, such as categories \u0026lsquo;social practices, rituals and festive events\u0026rsquo; and \u0026lsquo;performing arts\u0026rsquo; from the UNESCO domains of intangible cultural heritage. In other cases, we consolidated categories, for example, \u0026lsquo;food and feed\u0026rsquo; (NCP reporting categories), \u0026lsquo;food processing\u0026rsquo; (Ramsar Typology) and \u0026lsquo;gastronomy\u0026rsquo; (ethnoherpetological elements) to the category in our framework of \u0026lsquo;food and culinary heritage\u0026rsquo;. Where additional differentiation was feasible and logical, categories were further subdivided into finer subcategories to, as closely as possible, group together culturally informed interactions between people and freshwater animals, systematically describing similarities in the ways different cultural groups relate to species.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eDefining culture, cultural groups, and species\u003c/h3\u003e\n\u003cp\u003eWe focus on biocultural diversity, defined as \u0026ldquo;the range of human values and practices related to the dimensions of biodiversity at landscape, species and genetic levels\u0026rdquo;\u003csup\u003e\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e\u003c/sup\u003e, encompassing both biological and cultural diversity, and the interactions between the two that shape mutual coexistence. Our scope includes all peoples and is not limited to rural areas, as is sometimes the case in biocultural studies. We follow Reyes-Garcia et al. (2023)\u003csup\u003e\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u003c/sup\u003e, who define a cultural group as a community of practice who share a core set of beliefs, patterns of behaviour, and values. We identified cultural groups to the most specific level possible, based on literature provided, for example, \u0026lsquo;Tharu People from Chitwan district, Nepal\u0026rsquo; rather than \u0026lsquo;Indigenous People, Nepal\u0026rsquo; or \u0026lsquo;Tharu People\u0026rsquo;. Where reports from the literature provided only broad descriptors of sociocultural identity (e.g. Native Americans), we used these descriptors.\u003c/p\u003e \u003cp\u003eWe used the level of \u0026lsquo;species\u0026rsquo; (using scientific taxonomic nomenclature) as the unit of analysis for the biological diversity component in our framework, as this is the most widely used and recognised unit in conservation science and management\u003csup\u003e\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e\u003c/sup\u003e, for instance, the IUCN Red List of Threatened Species\u003csup\u003e\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e\u003c/sup\u003e. People often classify wildlife using folk taxonomies, which may align but are not always synonymous with scientific taxonomy. In our approach, we tie the cultural roles of a scientific species to a particular \u0026lsquo;folk taxon\u0026rsquo; or \u0026lsquo;ethnospecies\u0026rsquo; \u003csup\u003e\u003cspan additionalcitationids=\"CR48\" citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e\u003c/sup\u003e which required matching biological species with ethnospecies. In most instances, this was straightforward, as the source provided the scientific name of the species, the corresponding folk or vernacular name, or an image of the species. For example, \u0026lsquo;pike\u0026rsquo; in the United Kingdom corresponds to \u003cem\u003eEsox lucius\u003c/em\u003e, and images of a pike cannot be mistaken for those of any other local species. Where we found a lack of alignment, due to a folk-taxon corresponding to more than one scientifically defined species, all potential corresponding species, or an entire genus or family, were listed as the scientific species. For example, in Europe, multiple languages commonly use a single name - such as sturgeon (English), St\u0026ouml;r (German) or есетра (Bulgarian) \u0026ndash; to refer to all sturgeon species, especially \u003cem\u003eAcipenser\u003c/em\u003e spp. and sometimes also \u003cem\u003eHuso huso.\u003c/em\u003e In another example, in popular culture, \u0026lsquo;crocodiles\u0026rsquo; is used to refer indiscriminately to all true crocodile species of the family Crocodylidae. Where there is a lack of information in the literature to differentiate between candidate species that may correspond to a folk taxon, all potential species were noted, along with the uncertainty in attribution. In some instances, we only included the folk or vernacular name, due to an overwhelming number of corresponding species. For example, for some of the rituals related to frogs there were well over 50 potential species to include. In these instances, we included a note to explain this.\u003c/p\u003e\n\u003ch3\u003eExample Collection\u003c/h3\u003e\n\u003cp\u003eWith a draft complete framework designed, we then iteratively tested out and refined how well the initial categories fared against a first set of collated examples that covered the full range of categories defined and major world regions. We collected case examples of relationships between humans and freshwater fauna that were portrayed as or could be interpreted as cultural. Relationships between humans and freshwater fauna included knowledge, practices, beliefs and uses. Examples were collated from peer-reviewed publications, grey literature, personal knowledge of the authors and other disciplinary specialists, field data from authors\u0026rsquo; own work, online museum collections, as well as through a snowballing approach (i.e., searching relevant references cited in the studies identified). For each example, we collected the most specific information available in the reference or record referring to species, sociocultural and/or ethnolinguistic groups, and a summary of the relationships and knowledge (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). We aimed to include a geographically, culturally, and taxonomically diverse set of examples and, therefore, actively sought out examples from continents where we initially lacked examples. We included examples from both native and alien species. We aimed for a minimum of three examples for each initial category (and at least one for a subcategory). With expansion of the dataset during adaptive testing of the framework (see below), we collected some 612 case examples on which we based the finalisation of the framework. As the focus of this study is outlining the framework, we then selected the most informative subset of these cases (n\u0026thinsp;=\u0026thinsp;232) to illustrate the framework for this paper, comprising a first knowledge base on which to continue to build (Supplementary Table S3). Many of the examples represent secondary sources of knowledge. A potential limitation in such instances is that different actors use different lenses to interpret and report knowledge. Wherever possible, therefore, we prioritised the sources closest to the original context. For instance, where multiple sources were available, we would favour those produced by the communities themselves, such as drawing on cosmogonies from official websites of First Nations or Aboriginal groups, rather than relying on colonial accounts.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eInformation collected for each example taxon of cultural significance\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"3\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaxonomic Information\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eInformation on Sociocultural Group\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eInformation on the nature of the relationship and the associated knowledge\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u0026bull; Family\u003c/p\u003e \u003cp\u003e\u0026bull; Genus\u003c/p\u003e \u003cp\u003e\u0026bull; Species\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026bull; Name or description of sociocultural group (e.g. ethnolinguistic group, societal group)\u003c/p\u003e \u003cp\u003e\u0026bull; Place of knowledge/relationship\u003c/p\u003e \u003cp\u003e\u0026bull; Country (or countries) of knowledge/relationship\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u0026bull; Folk name(s) of ethnospecies used by give sociocultural group\u003c/p\u003e \u003cp\u003e\u0026bull; Description or illustration of the relationship and/or knowledge\u003c/p\u003e \u003cp\u003e\u0026bull; Notes on how ethnospecies was related to scientific species (if not immediately clear and intuitive e.g. if there is under differentiation of species within a folk taxon)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e\n\u003ch3\u003eAssigning case examples to domains, categories and subcategories\u003c/h3\u003e\n\u003cp\u003eA specific definition was developed for each category and subcategory in the framework. Where possible, established definitions were drawn from the literature. For example, the category \u0026ldquo;Tourism\u0026rdquo; uses the definition of tourism from the United Nations Statistical Commission (2008)\u003csup\u003e\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e\u003c/sup\u003e. In most cases, a specific definition was developed, drawing on broader conceptualizations found in the existing literature. For example, the category \u0026lsquo;Worldviews and belief systems\u0026rsquo; is defined as \u0026ldquo;the species plays a key or specific role in a cultural worldview, which is the perceptual framework of a cultural group, in which worldviews are \u0026lsquo;socially constructed realities which humans use to frame perception and experience\u0026rsquo; and refers to the \u0026lsquo;psychological, cognitive and affective determinants of behaviour\u0026rsquo;\u003csup\u003e\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e51\u003c/span\u003e\u003c/sup\u003e. It includes beliefs, such as on cosmology, cosmogonies, the afterlife, goals, values, and ethics. Cultural worldviews may or may not be distinguishable from religious or spiritual beliefs, depending on the culture \u003csup\u003e\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e\u003c/sup\u003e.\u0026rdquo; Where the definition described a comparatively straightforward concept, we did not reference other literature as, for example, for the category \u0026ldquo;Representation in Music\u0026rdquo; which is defined as \u0026ldquo;the species is represented in and/or inspiration for the creation of music, such as songs, rhymes, chants, and instrumental music.\u0026rdquo;\u003c/p\u003e \u003cp\u003eWe assigned each example in our dataset to one or more categories or subcategories in the draft framework. Where examples did not fit comfortably within the framework structure, (sub)categories were adjusted or new (sub)categories were developed, through an iterative process. Where examples illustrated multiple facets of the relationship, each of the different facets were included in the different relevant categories and, where necessary, in multiple categories to ensure full representation of cultural significance. Framework refinement continued until all collected examples could be readily assigned to one or more categories. Through this process, we refined the categorization structure to ensure theoretical relevance for freshwater fauna, with reassessment of the specificity and distinctiveness of each category ensuring that the framework was both inclusive and analytically robust. Finally, we organised categories into groupings of thematically similar relationships, comprising seven overarching domains, which were named to reflect their internal categories (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Domains are thematically defined by the groupings of categories attributed to them. For example, the domain (7) thematically defined as \u0026lsquo;creative expression\u0026rsquo;, is made up of the individually defined categories of \u003cem\u003evisual arts\u003c/em\u003e (7.1), \u003cem\u003eperforming arts\u003c/em\u003e (7.2), \u003cem\u003elanguage arts\u003c/em\u003e (7.3) \u003cem\u003eand other forms of creative inspiration\u003c/em\u003e (7.4).\u003c/p\u003e \u003cp\u003eThe resultant framework was a hierarchical three-tier system, with broad domain subdivided into categories, which were divided into subcategories where necessary (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). The hierarchical organization was designed to capture increasing specificity in the nature of human-species relationships, while maintaining conceptual clarity and practical applicability across diverse cultural contexts. Our example dataset includes multiple examples for each domain, category and subcategory, from which we have selected those that best illustrate each of these framework elements. We also included some specific cultural group and species pairings in the results repeatedly, to illustrate how the relationship between a given species and culture can have multiple facets. Similarly, we included relationships of different cultures with the same species or taxon, to illustrate how one species can have multiple significance across multiple cultures.\u003c/p\u003e\n\u003ch3\u003ePositionality statement\u003c/h3\u003e\n\u003cp\u003eThis research aims to provide a framework for identifying and categorising types of human-freshwater fauna relationships across multiple cultures, with the goal of improving cross-cultural comparisons and understanding of these relationships. Our author team includes people from different nationalities, disciplinary backgrounds, and positionalities, and both Indigenous and non-Indigenous scholars. We acknowledge that our approach, and the framework construct itself, is largely influenced by Western epistemic traditions, and is driven by the aim to facilitate understanding of the biocultural importance of freshwater fauna to scholars, practitioners, policymakers, stakeholders and rightsholders. We hope to contribute to respectful, clearer cross-cultural communication and recognition of cultural significance as an integral and vital part of freshwater ecosystem management and conservation. We appreciate that Indigenous Peoples, and different cultural groups or communities, might conceptualise, categorise, and understand relationships very differently, and through different lenses these categories may not be universally applicable, meaningful, or appropriate. We recognise these limitations of the framework, as well as those considered in the below and offer it, not as a universal approach, but as one possible tool to build a coherent evidence base that supports cross-cultural dialogue across knowledge systems. With this framework, we seek to provide a pragmatic and respectful tool for recognising the plurality of human-freshwater fauna connections, and for making these relationships more visible and actionable in conservation and heritage contexts.\u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eConsiderations on use and application of the framework\u003c/h2\u003e \u003cp\u003eA comprehensive understanding of the cultural significance of species necessitates deep engagement with the complex and often nuanced cultural contexts in which human\u0026ndash;other-than-human relationships are embedded. These relationships are inherently context-specific and can be challenging to convey to those outside a cultural group\u003csup\u003e\u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e53\u003c/span\u003e\u003c/sup\u003e. The framework we propose offers a structured approach for exploring cultural significance. It is not intended as a substitute for participatory, inclusive methodologies that respect the rights and autonomy of cultural groups, particularly in conservation and related applications. It serves as a tool to aid in systematising knowledge, and as a starting point for exploring the diverse ways in which people relate to freshwater species. Similarly, assignment of categories can be challenging to those without a deep understanding of a culture. We sought to reduce this limitation as much as possible by dividing categories at the most easily \u0026lsquo;quantifiable\u0026rsquo; part of the relationship (i.e., the human-animal or human-representation interaction), rather than using a framework based on values. For example, recreational fishing may be valued for a range of culturally specific reasons such as fun, additional food, or relationship-building. However, we class it as recreational fishing based on the activity, rather than the underlying justification or values.\u003c/p\u003e \u003cp\u003eAs with all research involving people, there are important ethical considerations in the collation and mobilisation of data on the relationships between human and other elements of nature\u003csup\u003e\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e\u003c/sup\u003e. A careful, sensitive and respectful approach to knowledge is particularly important with Indigenous and other local community knowledge, and we highlight the importance of proper engagement in understanding cultural engagements with freshwater taxa\u003csup\u003e\u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e55\u003c/span\u003e\u003c/sup\u003e. For example, some knowledge and other aspects of relationships between species and a given culture are sacred and/or secret knowledge\u003csup\u003e\u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e56\u003c/span\u003e\u003c/sup\u003e, and maintaining the sacred or secret nature of such knowledge is paramount. Our framework provides one approach to discuss or reference a relationship without giving any specifics on the knowledge itself, with the categories providing a potential placeholder for such knowledge in databases and/or discussions. This needs to be done on a case-by-case basis, as in some circumstances, collection or categorisation of knowledge could be unethical or inappropriate. We urge all users of this framework to consider which data are included in a cautious, respectful and rights-based manner.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eThe framework (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e\u003cem\u003e)\u003c/em\u003e organises culturally significant relationships between people and freshwater fauna into seven overarching domains, representing thematic groupings which broadly capture the diversity of human-fauna interactions. Each domain is divided into multiple categories, and some categories are further subdivided into subcategories. The full set of categorized relationships is provided in Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. To facilitate interpretation, detailed definitions of each category and subcategory are provided in the Supporting Information (Table S2). To explain the framework, we use a selection of brief references to examples, all of which are further detailed in the Supporting Information (Table S3), alongside a larger set of explanatory examples (n\u0026thinsp;=\u0026thinsp;232).\u003c/p\u003e\n\u003ch3\u003eDomain 1: Consumptive Use\u003c/h3\u003e\n\u003cp\u003eThis domain encompasses human cultural interactions involving the consumptive use of the species, alive or dead, and related knowledge. A particularly knowledge rich theme, it includes most interactions typically considered \u0026lsquo;material\u0026rsquo;, \u0026lsquo;tangible\u0026rsquo;, or \u0026lsquo;provisioning\u0026rsquo; \u0026mdash;such as food, material, and medicinal uses\u003csup\u003e\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e\u003c/sup\u003e\u0026mdash; as well as \u0026lsquo;intangible\u0026rsquo; or \u0026lsquo;immaterial\u0026rsquo; aspects of such uses. For instance, the fish \u003cem\u003eTenualosa ilisha\u003c/em\u003e is widely eaten in South Asia and exemplifies multiple categories of this Domain. In Bangladesh, it is important for \u003cem\u003efood security\u003c/em\u003e (1.1.1) and \u003cem\u003enutritional security\u003c/em\u003e (1.1.2), accounting for over 12% of Bangladesh\u0026rsquo;s total fish production and 1% of national gross domestic product \u003csup\u003e\u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e57\u003c/span\u003e\u003c/sup\u003e. Called \u003cem\u003eIlish\u003c/em\u003e in Bengali, it is consumed in \u003cem\u003eritual and social practices\u003c/em\u003e (1.1.4), such as Bengali New Year. Bengali Hindus also offer the fish to deities in other festivals\u003csup\u003e\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e58\u003c/span\u003e\u003c/sup\u003e. In Sindh, Pakistan, it is a prized food called \u003cem\u003ePalla\u003c/em\u003e, which is offered to guests as a sign of honour and respect (1.1.3 and 1.1.4) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Consumptive relationships can also have negative effects, as is the case of the fish Aymara (\u003cem\u003eHoplias aimara\u003c/em\u003e) of French Guiana. While its consumption is a \u003cem\u003edelicacy\u003c/em\u003e (1.1.3) for many local communities (Griffith \u0026amp; Mangione, unpub. data), eating the fish poses a health risk due to mercury contamination\u003csup\u003e\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e59\u003c/span\u003e\u003c/sup\u003e. The species complex collectively known as the Chinese Giant Salamander (\u003cem\u003eAndrias\u003c/em\u003e spp.) is also a \u003cem\u003edelicacy\u003c/em\u003e, as well as status symbol food in mainstream urban Chinese culture\u003csup\u003e\u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e60\u003c/span\u003e\u003c/sup\u003e. The latter cultural connection, along with local consumption in communities across its range, has pushed the salamander close to extinction in the wild\u003csup\u003e\u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e60\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eThe category \u003cem\u003eknowledge related to capture\u003c/em\u003e (1.2) includes traditional practices that enable locating, capturing, and killing of a species. Tharu fishers in Chitwan Nepal employ a method of attracting eels by constructing underwater stone piles (Griffith, unpub. data), represented under the subcategory of \u003cem\u003etracking, locating and attracting knowledge\u003c/em\u003e (1.2.1). The knowledge of how to weave Māori eel traps (Hīnaki) to capture \u003cem\u003eAnguilla\u003c/em\u003e spp. or tuna in the Māori language, is \u003cem\u003ecapturing knowledge\u003c/em\u003e (1.2.2)\u003csup\u003e\u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e61\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eHealing and pharmacological uses also form a category within \u003cem\u003eConsumptive Use\u003c/em\u003e. Many species are used in \u003cem\u003etraditional healing systems\u003c/em\u003e (1.3.1). For example, red toads (\u003cem\u003eSchismaderma carens\u003c/em\u003e) are sold in traditional medicine markets of Johannesburg, South Africa\u003csup\u003e\u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e62\u003c/span\u003e\u003c/sup\u003e. Similarly, fisher families of the Tocantins River, Brazil, use the fat of the poraqu\u0026ecirc; eletrico (electric eel, \u003cem\u003eElectrophorus electricus\u003c/em\u003e) as medicine and the electric shocks to treat rheumatism\u003csup\u003e\u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e63\u003c/span\u003e\u003c/sup\u003e. Certain species are used in \u003cem\u003econventional medicine and research\u003c/em\u003e (1.3.2), like zebrafish (\u003cem\u003eDanio rerio\u003c/em\u003e), a prominent model organism for the international scientific community\u003csup\u003e\u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e64\u003c/span\u003e\u003c/sup\u003e. \u003cem\u003eOther pharmacological uses\u003c/em\u003e (1.1.3) include poisons, mind-altering drugs, and drugs for spiritual or cultural rituals. For example, AmaZulu and AmaXhosa peoples of South Africa use nwenya, Nile crocodile (\u003cem\u003eCrocodylus niloticus\u003c/em\u003e), to make poisons and potions for strength\u003csup\u003e\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e65\u003c/span\u003e\u003c/sup\u003e, while Mats\u0026eacute;s hunters, Peru, use the Sapo (derived from secretions of the frog dowkiet, \u003cem\u003ePhyllomedusa bicolor)\u003c/em\u003e to enhance hunting success\u003csup\u003e\u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e66\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eC\u003cem\u003eonsumptive use\u003c/em\u003e also includes a category that captures the use of the animal or its parts in \u003cem\u003ecrafts, artefacts or tools\u003c/em\u003e (1.4.1), or items to be worn, such as \u003cem\u003eclothing or jewellery\u003c/em\u003e (1.4.2). Freshwater pearls from the cockscomb pearl mussel, 褶纹冠蚌 (\u003cem\u003eCristaria plicata\u003c/em\u003e), have been used for millennia in China, for ornamentation (1.4.2) \u003csup\u003e\u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e67\u003c/span\u003e\u003c/sup\u003e. In Medieval Western Europe, isinglass from sturgeon (\u003cem\u003eAcipenser\u003c/em\u003e spp. and \u003cem\u003eHuso huso)\u003c/em\u003e swim bladders was used as glue in book illumination\u003csup\u003e\u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e68\u003c/span\u003e\u003c/sup\u003e. Other categories include use for \u003cem\u003eenergy\u003c/em\u003e (1.5), such as oil from the eggs of the Arrau turtle (\u003cem\u003ePodocnemis expansa\u003c/em\u003e, Tartaruga) that Portuguese settlers used for lighting \u003csup\u003e\u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e69\u003c/span\u003e\u003c/sup\u003e. The name \u0026lsquo;candlefish\u0026rsquo; attributed to the Eulachon \u003cem\u003eThaeleichthy pacificus\u003c/em\u003e refers to the traditional practice of using the dried fish as a candle (1.5.1, \u003cem\u003eenergy from primary use\u003c/em\u003e), a method sometimes practiced by the Nisda\u0026rsquo;a First Nations people in Canada. They call the fish halmotkw, which can be translated as \u0026lsquo;salvation fish\u0026rsquo;, as the anadromous run of the fish was important for \u003cem\u003efood security\u003c/em\u003e (1.1.1), and today remains important for \u003cem\u003econsumption in ritual or social practices\u003c/em\u003e (1.1.4), especially the grease of the fish (1.1.6, \u003cem\u003efood processing\u003c/em\u003e)\u003csup\u003e\u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e70\u003c/span\u003e\u003c/sup\u003e. The use for \u003cem\u003ecompanionship and decoration\u003c/em\u003e (1.6) is also a category of this domain: 龙鱼, the Asian arowana (\u003cem\u003eScleopages formosus\u003c/em\u003e) is an exceptionally popular aquaria pet in Chinese cultures\u003csup\u003e\u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e71\u003c/span\u003e\u003c/sup\u003e. Finally, \u003cem\u003enon-use removal\u003c/em\u003e (1.8) refers to cases where a species is killed or removed without any intention of use, often due to a perceived negative cultural value, such as bad luck, as an ill omen, or belief in a negative interaction with another preferred species. For example, alligator gar (\u003cem\u003eAtractosteus spatula\u003c/em\u003e) have been viewed in recreational fishing communities in the USA as detrimental to \u0026lsquo;game fishes\u0026rsquo; and have been persecuted to the point of eradication in some areas\u003csup\u003e\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e72\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eDomain 2: Cultural Landscapes, Sites and Stewardship\u003c/h2\u003e \u003cp\u003eThis domain captures how the relationship between people and certain freshwater species shapes landscapes, including through the emergence of cultural landscapes, sites and places of settlement or worship, and stewardship practices (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). \u003cem\u003eSites and structures\u003c/em\u003e (2.1) refers to locations shaped through connections with specific species. We identify three time periods that indicate the origin of the \u003cem\u003esites and structures\u003c/em\u003e in their current (or final) location, aligned with the Ramsar typology of wetland-related activities\u003csup\u003e\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e\u003c/sup\u003e: \u003cem\u003eancient and prehistoric\u003c/em\u003e (pre-500CE); \u003cem\u003epost-classical\u003c/em\u003e (500-1599CE); and \u003cem\u003etraditional and modern\u003c/em\u003e (post-1599CE). \u003cem\u003eAncient and pre-historic sites and structures\u003c/em\u003e (2.1) include Mount Sandel, a Mesolithic Irish settlement site, likely sited due to reliance on eel, eascann (\u003cem\u003eAnguilla anguilla\u003c/em\u003e), and salmon, brad\u0026aacute;n (\u003cem\u003eSalmo salar)\u003c/em\u003e, from the River Bann\u003csup\u003e\u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e73\u003c/span\u003e\u003c/sup\u003e. Small-scale Karpfen (common carp, \u003cem\u003eCyprinius carpio\u003c/em\u003e) farming pond landscapes in Bavaria, Germany, have been maintained since the Middle Ages, and therefore fall under the subcategory of a \u003cem\u003epost-classical site\u003c/em\u003e (2.2). They are listed as Intangible Cultural Heritage by the German Commission for UNESCO (2023). The Chabdhi Barahi Temple in Tanahun, Nepal, established by the local Magar community in 1939, on instruction of the Goddess Barahi Devi, is a \u003cem\u003etraditional and modern site and structure\u003c/em\u003e (2.3). Sacred Rajabaam eels (\u003cem\u003eAnguilla bengalensis\u003c/em\u003e) in the temple pools are a key agent in worship (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), viewed variously as an incarnation, messenger, or protector of the goddess (Griffith \u0026amp; Basnet unpub. data). It is impermissible to fish for these eels locally. In Chile, a \u003cem\u003etraditional and modern site and structure\u003c/em\u003e (2.3) has arisen in Lake Llanquihue. In the 1970s, the industry based on non-native salmonids created a new ecosystem for the purpose of aquaculture and recreational fishing, now a multi-million dollar industry\u003csup\u003e\u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e74\u003c/span\u003e\u003c/sup\u003e, although the resultant declines in native species lead to conflicting views on the emergence of this new and novel ecosystem dominated by alien species\u003csup\u003e\u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e75\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003e \u003cem\u003eStewardship practices\u003c/em\u003e (2.2) are actions taken to protect, care for, or responsibly use the environment\u003csup\u003e\u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e76\u003c/span\u003e\u003c/sup\u003e, and reflect the cultural aspects of peoples\u0026rsquo; relationships with other elements of nature (Mattalia et al., 2025, in review). We follow the designation of Mattalia et al., (2025) in having three subcategories. \u003cem\u003eStewardship of target species\u003c/em\u003e (2.2.1) can be seen in the Solomon Islands, such as practices by Uluna and Sutahuri tribes in Guadalcanal. Members of these tribes enact stewardship of the eel mauvo or puleo (\u003cem\u003eAnguilla marmorata\u003c/em\u003e), refraining from harvesting them and strictly protecting them within their waterways (Hevalao unpub. data). Similarly, this species is an aquatic animal totem for Delailasekau village of the Naitasiri highlands of Fiji and therefore there is a taboo for killing and eating them across Fiji Islands (Rashni \u0026amp; Mailautoka, unpub. data). The Menominee Nation, USA, has a long history of stewardship of the culturally important n\u0026auml;m\u0026auml;'o or lake sturgeon (\u003cem\u003eAcipenser fulvescens\u003c/em\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), which is evident today in recovery of the species in the Winnebago System as a result of co-management by the Menominee and others\u003csup\u003e\u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e78\u003c/span\u003e\u003c/sup\u003e. The Yakel tribes of Tana islands, Vanuatu, maintain strong Kastom beliefs and protect sacred sites, including freshwater springs and river sources, demonstrating \u003cem\u003estewardship of an assemblage of species\u003c/em\u003e (2.2.2) such as \u003cem\u003eAnguilla\u003c/em\u003e spp. (eels) and \u003cem\u003eSicyopterus spp.\u003c/em\u003e (gobies). Water sources are often tied to ancestral spirits, with certain areas tabu (off-limits) for extraction or development of waterways (Hevalao unpub. data), which is \u003cem\u003estewardship of water and land\u003c/em\u003e (2.2.3). This can also be illustrated by the Nimiipuu People, whose water stewardship practices (affecting salmon) include knowledge of watershed protection and ongoing advocacy for breaching of the four lower Snake River dams that currently inhibit the free-flowing rivers required by native salmonids\u003csup\u003e\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e79\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eDomain 3: Knowledge Systems\u003c/h2\u003e \u003cp\u003eThis domain concerns the structure of knowledge systems and knowledge transmission across and within generations (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). While all categories in this framework represent types of knowledge (from knowledge of recipes to religion), this domain specifically focuses on the structure, storage, and transmission of knowledge. This may involve knowledge about the species themselves, or instances where the species play a role in transmitting other kinds of knowledge. Fundamental to the holding and transmission of species-related knowledge is its naming (3.1, \u003cem\u003enames\u003c/em\u003e): folk names and taxonomies are a culturally-rooted system of classification, conveying knowledge about the species and its cultural conception\u003csup\u003e\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e,\u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e80\u003c/span\u003e\u003c/sup\u003e. For example, \u003cem\u003eAnguilla anguilla\u003c/em\u003e is \u0026lsquo;eel\u0026rsquo; in the English language and English \u003cem\u003efolk taxonomies\u003c/em\u003e (3.1.1). The immature life stage during which eels move upriver from the sea is called \u0026lsquo;elver\u0026rsquo;, a corruption of the earlier \u0026lsquo;eelfare\u0026rsquo; meaning \u0026lsquo;passage of young eels upriver\u0026rsquo;. This name is part of the \u003cem\u003evocabulary of nature\u003c/em\u003e (3.1.2) as it transmits (or transmitted) knowledge about the species, as is the Japanese name, ito, for the Sakhalin taimen (Para\u003cem\u003ehucho perryi)\u003c/em\u003e, indicated in Kanji as 伊富. It is written using character parts for \"fish\" and \"ogre/demon\", reflecting its folkloric role as a monster in the water. The category \u003cem\u003ealternative word usage\u003c/em\u003e (3.1.3) includes instances when a taxon\u0026rsquo;s name also has other uses, such as \u0026lsquo;catfish\u0026rsquo; in modern English: both the folk name of the Siluriformes and a term for online identity deception\u003csup\u003e\u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e81\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eThe category \u003cem\u003eobservations\u003c/em\u003e (3.2) includes observations of species behaviour, life history, or ecology that are transmitted or interpreted culturally, such as through traditional or Indigenous knowledge. Some South African Indigenous Peoples consider bullfrogs (\u003cem\u003ePyxicephalus\u003c/em\u003e spp.) to rain from the sky, since these frogs spend most of the year aestivating underground, unseen until they emerge during heavy rains at the height of the rainy season\u003csup\u003e\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e\u003c/sup\u003e. Species can also be important in \u003cem\u003eknowledge acquisition\u003c/em\u003e (3.3) such the African clawed frog (\u003cem\u003eXenopus laevis\u003c/em\u003e), an important model organism for biology education\u003csup\u003e\u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e82\u003c/span\u003e\u003c/sup\u003e, thereby playing a key role in \u003cem\u003eknowledge transmission\u003c/em\u003e (3.4). Knowledge transmission can take many forms, such as in the S\u0026atilde;o Benedito community, Ilha do Ituqui, Brazil, where fishing for pirarucu (\u003cem\u003eArapaima gigas\u003c/em\u003e) is associated with various knowledges, and the practice of pirarucu fishing enables personal and social transfer or development of these knowledges\u003csup\u003e\u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e83\u003c/span\u003e,\u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e84\u003c/span\u003e\u003c/sup\u003e. A similar role is conferred through the importance of salmon (\u003cem\u003eOncorhynchus\u003c/em\u003e spp.) to the Nimiipuu People, for whom \u0026ldquo;the annual return of the salmon allows the transfer of traditional values from generation to generation\u0026rdquo; (Columbia River Inter-Tribal Fish Commission, 2006, p. 4)\u003csup\u003e\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e79\u003c/span\u003e\u003c/sup\u003e. Species can play a role in \u003cem\u003eknowledge inspiration\u003c/em\u003e (3.5), such as the electric eel (\u003cem\u003eElectrophorus electricus\u003c/em\u003e), a key inspiration for the development of batteries by the European scientific community\u003csup\u003e\u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e85\u003c/span\u003e\u003c/sup\u003e. They learnt of this knowledge from \u003cem\u003eobservations\u003c/em\u003e (3.2) from local fishers near Calabozo, modern day Venezuela, likely Indigenous Peoples.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eDomain 4: Worldviews, Belief Systems, and Cultural Identities\u003c/h2\u003e \u003cp\u003eThis domain includes human-freshwater fauna interactions that reflect a species\u0026rsquo; role in worldviews, belief systems, and identity formation (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). This includes species inclusion in socially constructed frameworks, such as religious or spiritual beliefs, and the oral or written traditions that support them, including myths, legends, and histories. This domain also includes the contribution of species to cultural or regional identity. These roles are often deeply embedded in how cultures make meaning, relate to the environment, and maintain a sense of belonging. For example, B\u0026auml;ru, the saltwater crocodile (\u003cem\u003eCrocodylus porosus\u003c/em\u003e), is considered protector of the Gumatj clan of the Yolŋu People of Arnhem Land, Australia\u003csup\u003e\u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e86\u003c/span\u003e\u003c/sup\u003e, a part of the Yolŋu \u003cem\u003eworldview and belief system\u003c/em\u003e (4.1). Also in Australia, pondi, the Murray cod (\u003cem\u003eMaccullochella peelii\u003c/em\u003e), is an important component of the Ngarrindjeri \u003cem\u003eworldview\u003c/em\u003e (4.1), including the Ngarrindjeri philosophy of being (known as Ruwel Ruwar)\u003csup\u003e\u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e87\u003c/span\u003e\u003c/sup\u003e. Pondi is essential in the origin of the Murray River and in the Ngarrindje \u003cem\u003enarrative account\u003c/em\u003e (4.3) of the Creation (Kaldowinyeri). A \u003cem\u003eworldview and belief system\u003c/em\u003e (4.1) of the Nguni People in South Africa is that frogs, specifically their croaking, is a sign that the ancestors approve of a ritual that was performed at the river\u003csup\u003e\u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e88\u003c/span\u003e\u003c/sup\u003e, and in Fiji, when a totem species appears out of the ordinary in freshwaters, it is seen as a good omen to the tribal landowners (Rashni \u0026amp; Mailautoka, unpub. data). Many species have a role in \u003cem\u003espirituality and religion\u003c/em\u003e (4.2), such as taimen (\u003cem\u003eHucho taimen\u003c/em\u003e), believed in Mongolia to be children of the river god, linking people and the spirit of the land and water. The role of species in worldviews, religion and narrative accounts is not always positive. The Chinese Giant Salamander has a negative cultural value in some Chinese communities, associated with bad luck and especially dead children (salamanders being transformed dead babies, or eating them), leading to its common name \u0026lsquo;baby fish\u0026rsquo; or 娃娃鱼\u003csup\u003e\u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e89\u003c/span\u003e\u003c/sup\u003e. For many communities in Amazonia, the river dolphin boto (\u003cem\u003eInia\u003c/em\u003e spp.) is strongly associated with enchantment and shapeshifting (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), often blamed for illegitimate pregnancies, the \u0026lsquo;children of the boto\u0026rsquo;. While the associated \u003cem\u003enarrative accounts\u003c/em\u003e (4.3) are oftentimes taken light-heartedly, they have also been used to mask violence against women\u003csup\u003e\u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e90\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eThis domain captures aspects of cultural or regional \u003cem\u003eidentity and relationships\u003c/em\u003e (category 4.4). For example, lafaek, the saltwater crocodile features in the Timorese belief system, lulik, in Timor-Leste, where there is \u003cem\u003ekinship\u003c/em\u003e (4.4.1) between certain cultural groups and the \u0026lsquo;ancestor crocodile\u0026rsquo; or Avo Lafaek, the grandfather crocodile \u003csup\u003e\u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e91\u003c/span\u003e\u003c/sup\u003e. Koena, the Nile crocodile, is the totem of the Bakoena clans of Lesotho and South Africa\u003csup\u003e\u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e92\u003c/span\u003e,\u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e93\u003c/span\u003e\u003c/sup\u003e, therefore playing a role in \u003cem\u003esupporting cultural identities\u003c/em\u003e (4.4.2). This even for the Bakoena who have moved to Lesotho, where crocodiles are not found. The crocodile is further a symbol of Lesotho\u0026rsquo;s national identity, as it features in the country\u0026rsquo;s coat of arms. Such relationships also provide terms for \u003cem\u003elandscape, sense of place, and connection to place\u003c/em\u003e (4.4.3): kwena is the name for \u003cem\u003eCrocodylus niloticus\u003c/em\u003e for the BaPedi People and is the place name for Mogalakwena municipality, and the Mogalakwena river, in South Africa\u0026rsquo;s Limpopo Province. Mogalakwena is a compound word referring to the might of the crocodile\u003csup\u003e\u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e94\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eThe domain also encompasses \u003cem\u003esymbolic or associative meaning\u003c/em\u003e (4.5). Zulu people in South Africa use the general term for frogs as an idiomatic expression (\u003cem\u003efigurative language\u003c/em\u003e, 4.5.2) for nudity\u003csup\u003e\u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e95\u003c/span\u003e\u003c/sup\u003e, threats of violence, and being honest\u003csup\u003e\u003cspan citationid=\"CR96\" class=\"CitationRef\"\u003e96\u003c/span\u003e\u003c/sup\u003e. The crocodile or kwena to the Bafokeng clan of the BaTswana people, also has \u003cem\u003esymbolic or associative meaning\u003c/em\u003e (4.5.1) as a symbol of peace, in addition to being their totem. The phrase \u0026ldquo;let the crocodile return to water\u0026rdquo; is used as a declaration of peace\u003csup\u003e\u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e97\u003c/span\u003e\u003c/sup\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), an \u003cem\u003eidiom, proverb, or turn of phrase\u003c/em\u003e (4.5.2). In Japan, the catfish, namazu 鯰 (primarily \u003cem\u003eSilurus asotus\u003c/em\u003e), is symbolic of earthquakes (4.5.1) and is used popularly and widely e.g., signs for roads closed after earthquakes. This can be explained by the \u003cem\u003eworldview\u003c/em\u003e (4.1), popularly known although no longer literally believed that earthquakes are caused by the thrashing of a giant catfish living under the islands of Japan\u003csup\u003e\u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e98\u003c/span\u003e\u003c/sup\u003e. Species names can also be used to evoke meaning through \u003cem\u003eassociative naming\u003c/em\u003e (4.5.3). For example, the term \u0026lsquo;buaya darat\u0026rsquo;, meaning \u0026lsquo;land crocodile\u0026rsquo;, is used to describe men who are playboys in Indonesia or Malaysia, evoking the predatory nature of crocodiles.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eDomain 5: Cultural and Social Practices\u003c/h2\u003e \u003cp\u003eThis domain contains categories that highlight how species are agents in shaping and maintaining cultures. It covers the involvement of fauna in social practices, ranging from everyday activities to celebratory rituals, social organisation, including gender roles and social stratification, and the rules or norms that influence relationships (such as laws, taboos, and resource use regulations). For example, the Amur carp (\u003cem\u003eCyprinus rubrofuscus\u003c/em\u003e), symbolic of determination and overcoming obstacles in Japan (4.4.4), is a key symbol in children\u0026rsquo;s day (previously boy\u0026rsquo;s day) in Japan. It is depicted in koinobori (carp streamers) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), part of category \u003cem\u003eceremonies, rituals, celebrations, and social practices\u003c/em\u003e (5.1). In the S\u0026atilde;o Benedito community, Brazil, hunting of the pirarucu (\u003cem\u003eArapaima gigas\u003c/em\u003e) successfully gives fishers prestige in their communities\u003csup\u003e\u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e84\u003c/span\u003e\u003c/sup\u003e, an example of the relationship with a species contributing to \u003cem\u003esocial organisation\u003c/em\u003e (5.2). Fisheries, such as that in Tono Reservoir in Ghana, often lead to gendered roles in fisher communities\u003csup\u003e\u003cspan citationid=\"CR99\" class=\"CitationRef\"\u003e99\u003c/span\u003e\u003c/sup\u003e, another example of \u003cem\u003esocial organisation\u003c/em\u003e (5.2). Such gendered interactions with harvested freshwater species can have profound social impacts, including negative impacts, such as with Nile perch (\u003cem\u003eLates niloticus\u003c/em\u003e) in Lake Victoria communities. The migration of Nile perch leads fishers (typically men) to follow the species, increasing the likelihood of engaging in high-risk sexual activities while away from their spouses. At the same time, declining catches in the lake\u0026mdash;particularly of native species\u0026mdash;have led some fishmongers (typically women) to form sexual relationships with fishers to secure fish. Together, these factors have contributed to increased human immunodeficiency virus prevalence in fishing communities\u003csup\u003e\u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e100\u003c/span\u003e\u003c/sup\u003e. This domain also encompasses \u003cem\u003erules or customary norms\u003c/em\u003e (5.3), such as the well-known and popularly referenced law in the United Kingdom that the monarch owns all unmarked mute swans in open water in the country (Exchequer of Pleas 1592), or the rule that killing ngweṋam, the Nile crocodile, is taboo for the Vhavenḓa people\u003csup\u003e\u003cspan citationid=\"CR101\" class=\"CitationRef\"\u003e101\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eDomain 6: Recreation, Leisure and Tourism\u003c/h2\u003e \u003cp\u003eThe \u003cem\u003erecreation, leisure and tourism\u003c/em\u003e domain encompasses interactions with a species \u0026ndash; non-consumptive, consumptive, and also representational (i.e., when the species itself is not necessarily physically involved) \u0026ndash; that are for the purposes of recreation, leisure, or tourism. For example, the Hecht or pike, \u003cem\u003eEsox lucius\u003c/em\u003e, is the focus of recreational fishing (6.1) in R\u0026uuml;gen, Germany, which includes both the consumption of the species and the capture of large \u0026lsquo;trophy\u0026rsquo; individuals\u003csup\u003e\u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e102\u003c/span\u003e\u003c/sup\u003e. Its local cultural role is reflected, for example, in a playground where the see-saw is shaped and painted as a pike, a representation of the species in sports and games (6.2.2). Such representation can even be purely conceptual, such as the idea of a crocodile (\u003cem\u003eCrocodylus\u003c/em\u003e spp.) in the children\u0026rsquo;s game \u0026lsquo;Please Mr Crocodile\u0026rsquo; played in various English-speaking cultures. This role of the species in traditional sports and games (6.2) can also involve the live animal (6.2.1), such as the fighting of siamese fighting fish ปลากัด, (\u003cem\u003eBetta splendens)\u003c/em\u003e in Thailand\u003csup\u003e103\u003c/sup\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Recreational practices can also include tourism (6.4.), such as cultural tourism (6.4.1), to visit the Chabdi barahi Temple and resident eels, in Nepal, or nature-based tourism (6.4.2) in Peru, where giant otter (\u003cem\u003ePteronura brasiliensis\u003c/em\u003e) sighting is a focus\u003csup\u003e104\u003c/sup\u003e. Species can also contribute to \u003cem\u003eaesthetics\u003c/em\u003e (6.3), such as dragonflies (\u003cem\u003eAeshnidae\u003c/em\u003e and \u003cem\u003eLibellulidae\u003c/em\u003e spp.) being appreciated for their appealing aesthetics in urban London, UK\u003csup\u003e105\u003c/sup\u003e.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003eDomain 7: Creative Expression\u003c/h2\u003e \u003cp\u003eThe final framework domain captures the expression of representations of the species in various creative aspects: visual, performing, and language arts. Such expressions are very often a reflection of the significance of species in the other six domains, as artists choose to use the arts to express different roles of species in their culture. For example, the saltwater and freshwater crocodiles of Australia (\u003cem\u003eCrocodylus porosus\u003c/em\u003e and C. \u003cem\u003ejohnstoni\u003c/em\u003e), b\u0026auml;ru and ng\u0026auml;w, are figures in the \u003cem\u003eartworks\u003c/em\u003e (7.1.1) of the Yolŋu people of Australia, with stylistic choices differentiating these culturally important species\u003csup\u003e\u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e86\u003c/span\u003e\u003c/sup\u003e. Knowledge of the two species is also conveyed through songs (\u003cem\u003emusic\u003c/em\u003e, 7.2.1.) and \u003cem\u003edance\u003c/em\u003e (7.2.2), where b\u0026auml;ru is depicted with arms extended in front of the body and ng\u0026auml;w by arms tucked in beside the body\u003csup\u003e\u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e86\u003c/span\u003e\u003c/sup\u003e. Folksongs of South Africa\u0026rsquo;s Venda and Tsonga cultures include frogs in lyrics\u003csup\u003e\u003cspan citationid=\"CR106\" class=\"CitationRef\"\u003e106\u003c/span\u003e,\u003cspan citationid=\"CR107\" class=\"CitationRef\"\u003e107\u003c/span\u003e\u003c/sup\u003e. This is taken further in Pedi culture by mimicking the frogs croak, while singing those folks songs. \u003cem\u003eVisual arts\u003c/em\u003e (7.1) include representations of the species in multiple respects, for example in crafts, artefacts, and tools (7.1.3), such as a West African Akan goldweight shaped like the eel, \u003cem\u003eAnguilla mossambica\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e) (Sheales 2014). In Betawi communities in Indonesia, the saltwater crocodile is represented in \u003cem\u003eculinary preparations\u003c/em\u003e (7.1.5) of Roti Buaya \u0026lsquo;bread crocodile\u0026rsquo;. The bread is symbolic of the commitment of the groom to act like the crocodile, who is believed by Betawai people to be faithful to its partner, and protect their children and partner from harm\u003csup\u003e\u003cspan citationid=\"CR108\" class=\"CitationRef\"\u003e108\u003c/span\u003e\u003c/sup\u003e. In Finland, thousands of years of cultural importance of the hauki or pike (\u003cem\u003eEsox lucius\u003c/em\u003e)(Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) can be seen through: prehistoric cave \u003cem\u003eartworks\u003c/em\u003e (7.1.1.) of the species acting as spirit guide to a shaman\u003csup\u003e\u003cspan citationid=\"CR109\" class=\"CitationRef\"\u003e109\u003c/span\u003e\u003c/sup\u003e; through important roles of the species in the Finnish national Epic Kalevala, a compilation of epic \u003cem\u003epoetry\u003c/em\u003e (7.3.2) and one of the most significant works of Finnish \u003cem\u003eliterature\u003c/em\u003e (7.3.1); through to modern popularity in recreational fishing and as food, reflected in artwork \u0026lsquo;The pike is a fish\u0026rsquo; by artist Outi Piiroinen in her series \u0026lsquo;Ode to Finnish food fish\u0026rsquo;\u003csup\u003e\u003cspan citationid=\"CR110\" class=\"CitationRef\"\u003e110\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003eOverlap in domains, and differing or conflicting relationships\u003c/h2\u003e \u003cp\u003eMany of the species used as examples are culturally significant in specific cultures, with relevance spanning multiple domains. For example, the \u003cem\u003econsumption of a species in ritual or social practices\u003c/em\u003e (\u003cem\u003econsumptive use\u003c/em\u003e, 1.1.4) evidently overlaps with \u003cem\u003esocial practices, rituals, and festive events\u003c/em\u003e (\u003cem\u003ecultural and social practices\u003c/em\u003e, 5.1), such as eating the Ilish fish on the first day of the Bengali year (1.1.4 and 5.1)\u003csup\u003e\u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e57\u003c/span\u003e\u003c/sup\u003e. Other categories and domains also overlap in some cases. For example, the use of the Siamese fighting fish in Thailand (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), part of the domain \u003cem\u003erecreation, leisure and tourism\u003c/em\u003e, has led to the species being popular in \u003cem\u003ecompanionship\u003c/em\u003e (1.6) as well as in \u003cem\u003esupporting cultural identities\u003c/em\u003e (4.4.2 in \u003cem\u003eworldviews, beliefs and identities\u003c/em\u003e) as the National Aquatic Animal of Thailand\u003csup\u003e\u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e103\u003c/span\u003e\u003c/sup\u003e. The species was inspiration for the national costume of Miss Universe Thailand in 2020, a role captured in the domain \u003cem\u003ecreative expression\u003c/em\u003e subcategories \u003cem\u003erepresentation in costume, clothing, and jewellery\u003c/em\u003e (7.1.4) and \u003cem\u003eother performing arts\u003c/em\u003e (7.2.5).\u003c/p\u003e \u003cp\u003eWithin some cultures, a given animal is immensely important across multiple domains, categories, and subcategories. This can be illustrated by considering some of the roles of Pacific salmon, chep, \u003cem\u003eOncorhynchus\u003c/em\u003e spp. in the culture of the Ainu of Hokkaido, Japan. The first salmon of the season is received in a ceremony known as Asircepnomi or Kamuycepnomi (1.1.3)\u003csup\u003e\u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e111\u003c/span\u003e\u003c/sup\u003e. This is a ceremonial \u003cem\u003eritual\u003c/em\u003e (5.1), which falls in the domain of \u003cem\u003ecultural and social practices\u003c/em\u003e, which also includes the \u003cem\u003ecustomary rule\u003c/em\u003e (5.3) that salmon should be killed with a specific ceremonial stick \u0026lsquo;isapa-kik-ni\u0026rsquo;\u003csup\u003e\u003cspan citationid=\"CR112\" class=\"CitationRef\"\u003e112\u003c/span\u003e\u003c/sup\u003e. This knowledge on how to correctly kill salmon, according to Ainu culture, also falls within the \u003cem\u003econsumptive use\u003c/em\u003e category \u003cem\u003eknowledge related to capture\u003c/em\u003e (1.2), specifically \u003cem\u003ekilling knowledge\u003c/em\u003e (1.2.3). This knowledge is conveyed between generations using songs\u003csup\u003e\u003cspan citationid=\"CR113\" class=\"CitationRef\"\u003e113\u003c/span\u003e\u003c/sup\u003e, a form of \u003cem\u003eknowledge transmission\u003c/em\u003e (3.4) within the domain \u003cem\u003eknowledge systems and transfer\u003c/em\u003e, but also \u003cem\u003emusic\u003c/em\u003e (7.2.1) a category within the \u003cem\u003ecreative expression\u003c/em\u003e domain. The songs convey a \u003cem\u003enarrative account\u003c/em\u003e (4.3), which falls within the domain of \u003cem\u003eworldviews, belief systems, and identities\u003c/em\u003e. In the narrative, Ainu people kill fish (Pacific salmon) with rotten sticks and, so, the fish god no longer sent the fishes to the Ainu: this would historically have jeopardised \u003cem\u003efood security\u003c/em\u003e (1.1.1). The first salmon ceremony is now practiced again despite many decades of criminalisation, and there is ongoing activism to restore Ainu rights to traditional salmon fishing beyond these ceremonies\u003csup\u003e\u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e111\u003c/span\u003e\u003c/sup\u003e. The role of salmon in contemporary movements for Ainu rights and cultural resurgence highlights the importance of salmon fishing as part of Ainu \u003cem\u003ecultural identity\u003c/em\u003e (4.4.2.). However, the salmon also have \u003cem\u003esymbolic meaning\u003c/em\u003e (4.4.4) through their connection to Indigenous People\u0026rsquo;s rights\u003csup\u003e\u003cspan citationid=\"CR114\" class=\"CitationRef\"\u003e114\u003c/span\u003e\u003c/sup\u003e. There are many additional facets of salmon significance, such as the use of salmon skins in shoes (1.4.2)\u003csup\u003e\u003cspan citationid=\"CR115\" class=\"CitationRef\"\u003e115\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eSome species have different roles in different communities, which can lead to conflict, particularly when consumption (especially overconsumption) of a species might jeopardise roles in other communities. For example, the diamondback terrapin (\u003cem\u003eMalaclemys terrapin\u003c/em\u003e) is Maryland\u0026rsquo;s state reptile and mascot for the University of Maryland (\u003cem\u003esupporting cultural identities\u003c/em\u003e 4.4.2.). However, this role was threatened when the species was heavily consumed as a \u003cem\u003edelicacy\u003c/em\u003e (1.1.3.) up until the 1990s, leading to its overexploitation and heightened risk of extinction; despite prohibition, the species was still being harvested as recently as 2016\u003csup\u003e116,117\u003c/sup\u003e. Such conflicting interactions can lead to profound cultural losses, such as the historical extirpation of the nam\u0026auml;'o or lake sturgeon in the Menominee Indian Reservation. Nam\u0026auml;\u0026rsquo;o has a key role in \u003cem\u003ecultural identity\u003c/em\u003e (4.4.2) including in the Metawin religion of the Menominee (\u003cem\u003espiritual or religious beliefs\u003c/em\u003e 4.2), in the creation story (\u003cem\u003enarrative accounts\u003c/em\u003e 4.3), as a clan totem (\u003cem\u003ekinship\u003c/em\u003e 4.4.1), as protector of rice beds, and as the tribal historian, keeping stories throughout its long life (\u003cem\u003eworldviews and belief systems\u003c/em\u003e 4.1)\u003csup\u003e\u003cspan citationid=\"CR118\" class=\"CitationRef\"\u003e118\u003c/span\u003e,\u003cspan citationid=\"CR119\" class=\"CitationRef\"\u003e119\u003c/span\u003e\u003c/sup\u003e. Throughout history the n\u0026auml;m\u0026auml;\u0026rsquo;o was important for Menominee \u003cem\u003efood and nutritional security\u003c/em\u003e (1.1.1,1.1.2), and \u003cem\u003etraditional medicines\u003c/em\u003e (1.3.1). The sturgeon was called by the sound of beating drums (an \u003cem\u003eobservation\u003c/em\u003e 3.2 of the sound \u0026lsquo;sturgeon thunder\u0026rsquo; hundreds of years before Western scientific documentation of the sound\u003csup\u003e\u003cspan citationid=\"CR120\" class=\"CitationRef\"\u003e120\u003c/span\u003e\u003c/sup\u003e), to Nam\u0026auml;'o Usk\u0026iacute;w\u0026auml;m\u0026icirc;t (meaning sturgeon spawning place, \u003cem\u003evocabulary of nature\u003c/em\u003e 3.1.2.). The first sturgeon is celebrated with a welcoming and feast in celebration and thanks for the arrival of the sturgeon (\u003cem\u003econsumption in ritual and social practices\u003c/em\u003e 1.1.4, \u003cem\u003esocial practices, ritual and festive events\u003c/em\u003e 5.1)\u003csup\u003e\u003cspan citationid=\"CR118\" class=\"CitationRef\"\u003e118\u003c/span\u003e,\u003cspan citationid=\"CR119\" class=\"CitationRef\"\u003e119\u003c/span\u003e\u003c/sup\u003e. However, commercial fishers from US settler communities historically viewed the lake sturgeon as a nuisance species, and therefore many sturgeons were persecuted through \u003cem\u003enon-use removal\u003c/em\u003e (1.8) or possibly for use in steamship boilers (\u003cem\u003eenergy\u003c/em\u003e, 1.5.1)\u003csup\u003e\u003cspan citationid=\"CR121\" class=\"CitationRef\"\u003e121\u003c/span\u003e\u003c/sup\u003e. Latterly, settler communities extensively overconsumed the species when it became valuable for its smoked meat and caviar (\u003cem\u003edelicacies\u003c/em\u003e 1.1.3) and isinglass (\u003cem\u003efood processing and preservation\u003c/em\u003e 1.1.6 and \u003cem\u003ematerial in crafts, artefacts or tools\u003c/em\u003e 1.4.1)\u003csup\u003e\u003cspan citationid=\"CR121\" class=\"CitationRef\"\u003e121\u003c/span\u003e\u003c/sup\u003e. Overexploitation and dam building led to collapses in sturgeon populations, and the extirpation of the n\u0026auml;m\u0026auml;'o in the Menominee Indian Reservation. This was a tragedy for the Menominee nation who had already had most of their land taken by settlers, and had been limited to the \u003cem\u003emodern site\u003c/em\u003e (2.1.3) of the Menominee Indian Reservation through a treaty in 1854. A major reason this land was chosen by the Menominee was it contained Nam\u0026auml;'o Usk\u0026iacute;w\u0026auml;m\u0026icirc;t\u003csup\u003e\u003cspan citationid=\"CR118\" class=\"CitationRef\"\u003e118\u003c/span\u003e\u003c/sup\u003e. This loss was described as \u0026ldquo;very cultural and spiritual loss, loss of food sustenance, loss of cultural identity\u0026rdquo; (David Nahwahquaw Grignon\u003csup\u003e\u003cspan citationid=\"CR119\" class=\"CitationRef\"\u003e119\u003c/span\u003e\u003c/sup\u003e). The importance of the n\u0026auml;m\u0026auml;'o was not forgotten, and stewardship by the Menominee Nation (\u003cem\u003estewardship of target species\u003c/em\u003e 2.2.1), including in collaboration with other groups such as \u003cem\u003erecreational fishers\u003c/em\u003e (6.1), has led to the restoration of sturgeon populations in the system, and the return of the species to Nam\u0026auml;'o Usk\u0026iacute;w\u0026auml;m\u0026icirc;t\u003csup\u003e\u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e78\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eOur novel framework provides a basis for systematically and hierarchically organising and analysing human-freshwater fauna interactions, structured into seven cohesive yet interconnected domains. It has the potential to serve applications in conservation policy, implementation and communication, sustainable management, interdisciplinary research, and the preservation of natural and cultural heritage. We provide a comprehensive lens for understanding a spectrum of human- fauna interactions that can be used to broaden utilitarian approaches to understanding and managing the natural world. This lens can help incorporate the relational values, social meanings, and intangible connections that are all too often missing from the discourse around freshwater biodiversity and ecosystem management.\u003c/p\u003e \u003cdiv id=\"Sec19\" class=\"Section2\"\u003e \u003ch2\u003eInsights into human-freshwater fauna relationships\u003c/h2\u003e \u003cp\u003eOur results show that, across cultures, there are multiple, complex and sometimes conflicting interactions between people and species of freshwater fauna, informing a broad range of very different cultural relationships in different communities. Such interactions range from consumptive uses to the role of species in framing worldviews. In this framework we moved away from binaries such as \u0026lsquo;material and immaterial\u0026rsquo; and \u0026lsquo;tangible and intangible\u0026rsquo;, as such dualisms can artificially segregate meaning and practice. We argue that the use of our framework provides room for a more comprehensive understanding between people and freshwater fauna. For example, the relationships between Sindhi people and the fish palla could be collapsed down to \u0026lsquo;provision of food\u0026rsquo; when using the ecosystem services framework. In contrast, our framework offers a method to resist such oversimplification: palla are important in foodways, and are sacred, they are pilgrims who must be able to undertake their pilgrimage (which is also their migration), they are favoured as an offering to the saint Khwaja Khizar\u003csup\u003e\u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003e122\u003c/span\u003e\u003c/sup\u003e, and are served as a sign of honour and respect to guests. The framework also offers a way to see how different versions of sacredness and respect for species are shown in different cultures: to some, a sacred species is one that should not be killed, such as the crocodile B\u0026auml;ru for the Yolŋu people in Australia\u003csup\u003e\u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e86\u003c/span\u003e\u003c/sup\u003e, to others, a sacred species is one that is used, but with respect and restraint for the gift of their lives, such as the lake sturgeon, nam\u0026auml;\u0026rsquo;o for the Menominee people in the USA\u003csup\u003e\u003cspan citationid=\"CR118\" class=\"CitationRef\"\u003e118\u003c/span\u003e\u003c/sup\u003e. Different worldviews shape different relationships with other-than-human nature, and each relationship can only be fully understood within its specific cultural context. However, our framework provides a valuable tool for recognising these interactions from both external and global perspectives. The framework supports cross-cultural communication and understanding of human-fauna interactions in freshwater ecosystems, whether they occur in societies different from, or inclusive of, our own. Furthermore, the examples highlight how culture significance is sometimes tied directly to a single scientific species, and other times a broader range of taxa are referred to by a given ethnospecies name, and therefore cultural significance and the scientific species concept are not always congruent.\u003c/p\u003e \u003cp\u003eOur examples and framework illustrate that the impacts of interactions of people and freshwater species can be positive, negative or neutral for people and/or other components of nature. These positive and negative contributions can be seen as parts of nature\u0026rsquo;s contributions to people, one of the frameworks with which we have aligned our work\u003csup\u003e\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e\u003c/sup\u003e. For example, the relationship between Finnish people and pike, hauki, is one in which pike provides positive contributions to people: from foodways to recreational fishing, to roles in narratives including folktales and the national epic, a feature in art from prehistoric caves paintings to modern design, interactions with pike shape and enrich Finnish culture (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Conversely, some relationships result in negative contributions to people, for example the changes to the ecosystem and associated fisheries in Lake Victoria after Nile perch introduction, with the consumptive relationship with the species in combination with its role in social organisation lead to profound negative social and public health impacts\u003csup\u003e\u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e100\u003c/span\u003e\u003c/sup\u003e. Particular interactions may have different impacts on different people in a community, such as the beliefs and associated narratives of the boto encantado (supernatural river dolphins) in Amazonia being valued stories for exploring concepts of cultural change and identity, but also being used to mask sexual violence\u003csup\u003e\u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e90\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eOur results also highlight influence in the other direction, of \u0026lsquo;people\u0026rsquo;s contributions to nature\u0026rsquo;. Positive contributions of people to nature are perhaps most clearly seen in the category of stewardship practices (2.2) which follows Mattalia \u003cem\u003eet al.\u003c/em\u003e, (2025, in review), but closely aligns with cultural values of species that are captured in other domains. For example, the positive impact of the Uluna and Sutahuri tribes on the eel mauvo or puleo, who see eels as guardians of river health, and in turn enact stewardship practices such as strictly protecting them, or the Nimiipuu people whose activism to push for breaching dams on the lower Snake river contributes to the stewardship of migratory salmonids\u003csup\u003e\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e79\u003c/span\u003e\u003c/sup\u003e. Such examples of positive contributions of people to nature are abundant among Indigenous Peoples, but are not limited to them. An example from recreational fishers in Lower Saxony, Germany, is a transdisciplinary and co-developed project which investigated how different stewardship practices (targeted stocking vs whole ecosystem restoration) could enhance populations of fishes important to angling clubs and their members\u003csup\u003e\u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e123\u003c/span\u003e\u003c/sup\u003e. Negative contributions of people to nature can arise from interactions that benefit humans at nature\u0026rsquo;s expense\u0026mdash;for example, the overconsumption of Chinese giant salamanders, has driven the species to near-extinction in the wild\u003csup\u003e\u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e60\u003c/span\u003e\u003c/sup\u003e. They can also stem from perceived negative interactions, such as non-use removal, which has led to local extirpations of alligator gar by U.S. recreational fishers who see the species as a competitor for prized game fish\u003csup\u003e\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e72\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eIn sum, the framework we present and its illustrative examples highlight that the relationships between people and freshwater fauna are multidirectional, with contributions of people to other species, and from other species to people. The framework defines interactions and relationships, rather than uses and contributions, and in so doing considers this multidirectionality. This aligns with more recent broadening of the NCP approach to not just provide for contributions of nature to people, but also \u0026lsquo;bidirectional relationships that include reciprocity\u0026rsquo;\u003csup\u003e\u003cspan citationid=\"CR124\" class=\"CitationRef\"\u003e124\u003c/span\u003e\u003c/sup\u003e, although the semantic lack of recognition of this in the term \u0026lsquo;nature\u0026rsquo;s contributions to people\u0026rsquo; has been criticised\u003csup\u003e\u003cspan citationid=\"CR125\" class=\"CitationRef\"\u003e125\u003c/span\u003e\u003c/sup\u003e. The concept of reciprocity in our relationships with nature, including with water bodies and the (other) lives they contain, is central to many Indigenous worldviews, and also increasingly recognised by others as central to sustainability transformations\u003csup\u003e\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e,\u003cspan additionalcitationids=\"CR127\" citationid=\"CR126\" class=\"CitationRef\"\u003e126\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR128\" class=\"CitationRef\"\u003e128\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec20\" class=\"Section2\"\u003e \u003ch2\u003eContribution of the framework to systematically organising and understanding different human freshwater-fauna interactions\u003c/h2\u003e \u003cp\u003eThe seven domains are broad thematic groupings that facilitate our understanding of complex relationships between humans and freshwater fauna, and the role of ethnospecies in shaping the connection between culture, interactions and the concept of scientific species. The domain groupings were emergent from the more specific, and individually defined, categories. In some cases, interactions could be distinguished at a finer, subcategory scale. For example, distinguishing between fishing for food security (1.1.1), delicacies (1.1.3), or ritual consumption (1.1.4), in the category food and culinary heritage (1.1), allows capturing differing roles in culinary heritage, spiritual practices, or as a staple food, a delicacy, or a culturally significant ingredient. For some categories, such as narrative accounts, we found that a finer granularity was not possible for a universal framework. This is because the categorization of stories \u0026ndash; such as whether they are seen to be histories, folktales, myths, or guidance \u0026ndash; varies between cultures. A culturally specific application of the framework could, however, develop culturally appropriate subcategories.\u003c/p\u003e \u003cp\u003eOur categories and subcategories are necessarily overlapping. Some subcategories always overlap with another category, such as examples placed in \u003cem\u003econsumption in ritual and social practices\u003c/em\u003e (1.1.4) always also falling in \u003cem\u003esocial practices, rituals and festival events\u003c/em\u003e (5.1). Similarly, interactions may fall in multiple domains and categories for a particular human-fauna relationship. We highlight this inevitable overlapping as important in recognising that relationships with fauna can be plural, multifaceted, and difficult to compartmentalize. The fighting of pla kat or Siamese fighting fish (\u003cem\u003eBetta splendens\u003c/em\u003e) is not simply a pet (consumptive use) or part of a game (recreation), but an important traditional cultural and social practice that is part of Thai national identity (worldviews and identity). Our finding that the relationship between a culture and species often includes many facets that fit into multiple distinct categories shows the plurality and complexity of considering the cultural significance of freshwater fauna. These examples highlight how many species have multiple and intersecting or amplifying importances to certain cultures.\u003c/p\u003e \u003cp\u003eIt is increasingly recognised that solutions to the global biodiversity crisis, most acutely exemplified in freshwater ecosystems, will require approaches that are transdisciplinary, weave multiple knowledges, accommodate diverse worldviews, and are Indigenous-led when in lands with Traditional Owners\u003csup\u003e\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e,\u003cspan citationid=\"CR126\" class=\"CitationRef\"\u003e126\u003c/span\u003e,\u003cspan citationid=\"CR129\" class=\"CitationRef\"\u003e129\u003c/span\u003e,\u003cspan citationid=\"CR130\" class=\"CitationRef\"\u003e130\u003c/span\u003e\u003c/sup\u003e. The framework presented here contributes to transdisciplinary approaches by connecting species (the unit typically used by natural scientists), to ethnospecies (the unit by which people conceptualise species), and to cultural significances in a systematic way that is conceptually simple to understand. The thematic and hierarchical structuring of the framework allows both an in-depth exploration of the multiple facets of the role of a given taxa in a particular culture as well as of the breadth of different types of importance between cultures, species, or globally. The multifaceted relationship between humans and eels (\u003cem\u003eAnguilla\u003c/em\u003e spp.) encompassing food security, religious and ritual practice, and cultural identity across the world, demonstrates how a single taxon can fulfil multiple cultural roles and situational meanings, and traverse political borders (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Additionally, the framework offers a way to approach documenting and understanding differing roles in different cultures, which may be contradictory or conflicting. For example, the overexploitation of lake sturgeon / nam\u0026auml;\u0026rsquo;o by US settlers due to non-use removal and later view of them as food delicacies, combined with a lack of stewardship in settler culture, conflicted with the highly important role of nam\u0026auml;\u0026rsquo;o in Menominee culture. This led to both negative impacts on the species and on a culture. Identification of contradictory relationships with the same species \u0026ndash; or species in the same ecosystem \u0026ndash; to different communities, is vital for understanding the conflicts that underscore much of biodiversity conservation and management of the natural world\u003csup\u003e\u003cspan citationid=\"CR131\" class=\"CitationRef\"\u003e131\u003c/span\u003e\u003c/sup\u003e. Of course, understanding of different interactions between people and fauna is just one part of conflict resolution, and occurs in the complex setting of trust, justice and power between the different communities involved\u003csup\u003e\u003cspan citationid=\"CR132\" class=\"CitationRef\"\u003e132\u003c/span\u003e\u003c/sup\u003e. This framework seeks to describe the interaction between people and freshwater animals, but we recognise that there are many values, importances, power imbalances, histories and human-human relationships that shape these interactions.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec21\" class=\"Section2\"\u003e \u003ch2\u003eApplications of the framework in interdisciplinary understanding\u003c/h2\u003e \u003cp\u003eThe differences in language and terminology used by different disciplines can make interdisciplinary work challenging. The framework helps provide a \u0026lsquo;translating lens\u0026rsquo; through which different types of scholarship can be understood more clearly, by providing a consistent framing vocabulary to the complexity of explaining culture. In the environmental humanities, for instance, approaches are often specified and relevant terminology used that natural scientists are unlikely to have come across, such as \u0026lsquo;posthumanism\u0026rsquo;, or \u0026lsquo;more-than-human\u0026rsquo;, but such work may contain in-depth discussions of human-species interactions relevant to a manager working with certain species. Similarly, fisheries scientists may write papers densely laden with terms such as \u0026lsquo;catch per unit effort\u0026rsquo; or \u0026lsquo;stock recruitment relationship\u0026rsquo; that may be alien to an environmental humanist. Both works may relate to the important role of a fish species in foodways and cultural identity, and therefore, the responsibility of communities to steward the species. This role could be outlined in a simple and clear way by applying our framework to these interactions, with reference to the more specific, disciplinary literature.\u003c/p\u003e \u003cp\u003eAdditionally, by categorizing significance from different cultures together, the framework can aid in cross-cultural understanding, as users can identify examples from their own culture, and use this to better recognize importance in other cultures. For example, the European eel is a culturally important food in the Netherlands (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e), but also a critically endangered species. For a Dutch person, understanding the cultural context of eating a critically endangered species (e.g., personal experiences of eating smoked eels with family) may help in building understanding and empathy as to why critically endangered species are also eaten in other cultures. Improving communication and understanding can also extend beyond preservation of biocultural diversity, into other areas, such as public health and medicine, which are also deeply embedded in cultural context. For example, small-scale gold mining leads to mercury contamination of water and then biomagnifies through trophic webs, creating public health and conservation concerns\u003csup\u003e\u003cspan citationid=\"CR133\" class=\"CitationRef\"\u003e133\u003c/span\u003e,\u003cspan citationid=\"CR134\" class=\"CitationRef\"\u003e134\u003c/span\u003e\u003c/sup\u003e. Dietary recommendations on which fish species are safe to consume \u0026ndash; or not \u0026ndash; through examining the nutritional, ecological, and market attributes, and mercury risks is a potential risk mitigation strategy\u003csup\u003e\u003cspan citationid=\"CR135\" class=\"CitationRef\"\u003e135\u003c/span\u003e\u003c/sup\u003e. A recent study shows that large, migratory fish in the Amazon (e.g. \u003cem\u003eBrachyplatystoma rousseauxii\u003c/em\u003e) have low nutrient density and high mercury content, and are also more threatened, and it suggested recommendations and messaging could discourage their consumption\u003csup\u003e\u003cspan citationid=\"CR135\" class=\"CitationRef\"\u003e135\u003c/span\u003e\u003c/sup\u003e. Understanding the cultural context, values and traditions around fish consumption, and integrating them with other information sources, could strengthen such an approach. In French Guiana certain species, including fish such as \u003cem\u003eB. rousseauxii, Hoplias aimara\u003c/em\u003e, and crocodylians such as \u003cem\u003ePaleosuchus trigonatus\u003c/em\u003e, have high mercury risks if consumed, but have a high cultural value for consumption (1.2.3), recreational hunting and fishing (6.1) and knowledge sharing of fishing techniques between generations (3.4), which can lead to continued consumption even in the knowledge of mercury risks.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec22\" class=\"Section2\"\u003e \u003ch2\u003eApplication and policy implications of the framework in biodiversity conservation\u003c/h2\u003e \u003cp\u003eThe framework proposed can complement ecology-oriented approaches for freshwater biodiversity conservation through different channels. Our biocultural approach highlights the bonds between freshwater fauna and various cultural groups in different regions and provides justifications for the protection of threatened freshwater fauna, beyond their ecological roles. Moreover, the biocultural framework can help gain momentum among the public for freshwater ecosystems to be experienced as vital parts of the environment upon which our society depends\u003csup\u003e\u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e136\u003c/span\u003e\u003c/sup\u003e. Freshwater ecosystems have often been regarded simply as a resource for clean water and food, with freshwater fauna such as fish, in many cases, not being considered wildlife, which could have negative impacts on the conservation of freshwater fauna\u003csup\u003e\u003cspan citationid=\"CR137\" class=\"CitationRef\"\u003e137\u003c/span\u003e,\u003cspan citationid=\"CR138\" class=\"CitationRef\"\u003e138\u003c/span\u003e\u003c/sup\u003e. By recognizing the cultural significance of freshwater fauna, researchers and conservation practitioners can work together to raise public awareness and increase their willingness to contribute to freshwater conservation through targeted communications that enhance emotional connections between human and freshwater fauna. Emotions, values and personal experience have been shown to be crucial for decision making, with emotionally driven storytelling demonstrably more effective than \u0026lsquo;fact-based\u0026rsquo; narratives in shifting attitudes\u003csup\u003e\u003cspan citationid=\"CR139\" class=\"CitationRef\"\u003e139\u003c/span\u003e\u003c/sup\u003e. As conservation research and practice remain insufficient in the face of current threats to freshwater biodiversity, freshwater conservationists widely acknowledge that losses can only be stemmed by a paradigmatic shift in relationships between people and freshwater biodiversity\u003csup\u003e\u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e136\u003c/span\u003e,\u003cspan citationid=\"CR140\" class=\"CitationRef\"\u003e140\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eGiven the widely observed shifting baseline syndrome for freshwater fauna, where each generation accepts a more degraded ecological state as the baseline\u003csup\u003e\u003cspan citationid=\"CR141\" class=\"CitationRef\"\u003e141\u003c/span\u003e,\u003cspan citationid=\"CR142\" class=\"CitationRef\"\u003e142\u003c/span\u003e\u003c/sup\u003e, highlighting past human-freshwater fauna links and revitalizing lost cultural traditions and stewardship practices holds the potential for engaging younger generations in freshwater conservation and supporting more ambitious goals for restoration. In some instances, traditions may have long been obscured, for example the once important and multifaceted role of the eel in British society\u003csup\u003e\u003cspan citationid=\"CR143\" class=\"CitationRef\"\u003e143\u003c/span\u003e\u003c/sup\u003e. In such cases, an approach to understanding historical or obscure significance using the framework we outline can identify emotive and engaging local or national examples, to garner interest and engagement in freshwater conservation. In other instances, this may be a matter of preserving and supporting the revitalization of cultural traditions and practices that are being lost due to colonisation, cultural suppression, or social and environmental change. For example, in parts of Guadalcanal, Solomon Islands, the Uluna and Sutahuri tribes revere freshwater eels as guardians of river health, enacting stewardship practices such as refraining from harvesting them and strictly protecting them within their rivers and streams. Unlike in other regions where eels are a food source, this tribe views them as essential for maintaining river health and cleanliness. The eel\u0026rsquo;s role in the ecosystem aligns with their spiritual beliefs, reinforcing non-extractive conservation practices. Similarly, in Vanuatu, traditional fishing taboos regulate when and how certain species can be caught, ensuring sustainable use. By revitalizing these stewardship practices, younger generations can develop a stronger connection to freshwater biodiversity, reinforcing conservation efforts in both customary and scientific contexts. Such actions can be part of a broader place-based biocultural approach. For example, in the Solomon Islands, place-based conservation efforts\u0026mdash;such as Locally Managed Marine Areas (LMMAs), and the iTina and Tina Rivers study which includes local rangers for boundary monitoring\u0026mdash;integrate traditional ecological knowledge with modern conservation techniques, ensuring that freshwater resources are protected through a mix of Indigenous and scientific strategies, aligning conservation with cultural narratives and ancestral responsibilities.\u003c/p\u003e \u003cdiv id=\"Sec23\" class=\"Section3\"\u003e \u003ch2\u003eFuture directions and next steps\u003c/h2\u003e \u003cp\u003eThis framework to categorise the cultural significance of freshwater fauna provides an approach to conceptualise and organise interactions between people and freshwater fauna, and consequently perhaps the most direct application is to use the framework as a tool for collecting and understanding data on these interactions. For example, the framework could be used to develop more comprehensive and standardised assessments of the implications of freshwater biodiversity loss, or potentials of restoration, or it could be incorporated into tools to measure conservation impact, such as the Open Standards for the Practice of Conservation of the Conservation Measures Partnership\u003csup\u003e\u003cspan citationid=\"CR144\" class=\"CitationRef\"\u003e144\u003c/span\u003e\u003c/sup\u003e. The framework could be used as a basis for knowledge or database collation on culturally informed interactions between people and freshwater species in particular basins or regions or for particular taxonomic groups. For example, migratory fishes have been identified as needing conservation for their ecological and socioeconomic value, with the concept of identifying \u0026lsquo;Global Swimways\u0026rsquo; proposed to address this need\u003csup\u003e145146\u003c/sup\u003e. However, so far, no social criteria have been suggested for Swimways, as they are seen as difficult to quantify\u003csup\u003e\u003cspan citationid=\"CR146\" class=\"CitationRef\"\u003e146\u003c/span\u003e\u003c/sup\u003e. The framework presented here provides a systematic method to approach this issue. Similarly, freshwater megafauna have been identified as a group in particular need of urgent conservation action\u003csup\u003e\u003cspan citationid=\"CR147\" class=\"CitationRef\"\u003e147\u003c/span\u003e\u003c/sup\u003e, and as having strong potential to serve as freshwater flagship species\u003csup\u003e\u003cspan citationid=\"CR148\" class=\"CitationRef\"\u003e148\u003c/span\u003e\u003c/sup\u003e, a role that is especially important given repeated calls to identify such species to help raise the profile of their threatened aquatic ecosystems\u003csup\u003e\u003cspan citationid=\"CR136\" class=\"CitationRef\"\u003e136\u003c/span\u003e,\u003cspan citationid=\"CR140\" class=\"CitationRef\"\u003e140\u003c/span\u003e\u003c/sup\u003e. However, flagship species operate as cultural assets, particularly through their presence in pre-existing cultural framings, alongside alignment with current cultural trends and political conditions\u003csup\u003e\u003cspan citationid=\"CR149\" class=\"CitationRef\"\u003e149\u003c/span\u003e\u003c/sup\u003e. This framework could provide a more effective approach to identifying potential flagship species, as well as for identifying with which communities a species may, or may not, galvanise action.\u003c/p\u003e \u003cp\u003eOur focus in this research has been on species of freshwater fauna, due to the particularly acute status of freshwater biodiversity, and the highly interrelated relationships between humans and all aspects of freshwaters and freshwater life. However, many of the species in our examples are migratory or multi-realm occupants, and spend part of their lives in marine systems. Given that relationships with such species do not stop at the coast, nor should consideration of their cultural roles in management or conservation. Investigations into how well the framework can be applied more broadly to all aquatic fauna, or to flora and funga, and beyond aquatic habitats to those on land, are interesting future avenues to be explored.\u003c/p\u003e \u003cp\u003eFinally, to borrow from Kimmerer\u0026rsquo;s (2012, p. 319)\u003csup\u003e\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u003c/sup\u003e thinking on teaching traditional ecological knowledge as a \u0026ldquo;prompt to the imagination, a mirror in which western society can examine its own relationship to the living world\u0026rdquo;, perhaps our framework can contribute to introducing and inspiring people who do not currently have reciprocal relationships with freshwater fauna to reconsider their own relationships and interactions. By presenting other ways of relating, and providing a way to systematically consider these as a complement to scientific thinking does not try to separate the tangible and intangible, our work could be a prompt to imagine new ways of relating to, and of conserving, freshwater species, such that these interactions are possible into the future.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e"},{"header":"Declarations","content":"\u003ch2\u003eAuthor Contributions\u003c/h2\u003e \u003cp\u003ePG, RET, FH, SK, PP and SCJ conceived the initial concept, which was initially developed by RET and PG with inputs from FH, SK, PP and SCJ. Example collection and contribution was conducted by PG, RET, FMP, LKFC, FH, RSH, SK, KKM, and BR. Framework definitions were developed by PG, RET and FMP, with extensive input from other authors. Framework refinement and completion was led by PG, with all authors contributing. PG took the lead in writing the manuscript, with all authors contributing sections, comments, or refinements to multiple drafts. All authors provided critical feedback and helped shape the research, analysis and manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgements\u003c/h2\u003e \u003cp\u003eThis work has been supported by the Leibniz Competition project \u0026lsquo;Freshwater Megafauna Futures\u0026rsquo; to SCJ and PG. RT recognises the research fellowship support of LE STUDIUM Loire Valley Institute for Advanced Studies, Orl\u0026eacute;ans and Tours, France, with funding from the European Union\u0026rsquo;s Horizon 2020 research and innovation programme under Marie Skłodowska-Curie grant agreement No. 665790, for a project on \u0026lsquo;Biological and cultural diversity, and flow regimes \u0026ndash; a unifying approach for managing riverscapes\u0026rsquo; which helped catalyse the collaboration reflected in this paper. RET further acknowledges funding support from The Nature Conservancy in framework drafting. FH acknowledges the support from the Chinese Academy of Sciences (E355S122). SK acknowledges the funding support from an anonymous donor to The Nature Conservancy. VRG acknowledges support from the \u0026ldquo;Mar\u0026iacute;a de Maeztu\u0026rdquo; Programme for Units of Excellence of the Spanish Ministry of Science and Innovation (CEX2019-000940-M). The authors would like to thank all the participants of a hybrid workshop on the \u0026lsquo;cultural importance of freshwater biodiversity\u0026rsquo; at the World Biodiversity Forum, 30 June, 2022, Davos, Switzerland, whose varied contributions helped further stimulate discussion around this topic. The authors would like to thank David Tickner and David Boseto for their helpful comments and suggestions on drafts of, or discussions about, the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eLinton, J. \u003cem\u003eWhat is water? The history and crisis of a modern abstraction\u003c/em\u003e. (BC:UBC Press, 2010).\u003c/li\u003e\n\u003cli\u003eMcCool, S. F., Clark, R. M., Stankey, G. H. \u0026amp; Mazza, R. Water, Recreation, and Society: Shifting Demands, Rising Concerns, Growing Complexity. in \u003cem\u003eWater and People: Challenges at the Interface of Symbolic and Utilitarian Values\u003c/em\u003e (eds. McCool, S. F., Clark, R. M. \u0026amp; Stankey, G. H.) 1\u0026ndash;18 (U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, 2008).\u003c/li\u003e\n\u003cli\u003eKummu, M., De Moel, H., Ward, P. J. \u0026amp; Varis, O. How Close Do We Live to Water? 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Soc.\u003c/em\u003e \u003cstrong\u003e13\u003c/strong\u003e, 95\u0026ndash;104 (2015).\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"","lastPublishedDoi":"10.21203/rs.3.rs-6830897/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6830897/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eFreshwater ecosystems support rich biological and cultural diversity, each enhanced through biocultural co-evolution. However freshwater species, and their relationships with people, face escalating pressures from global change, threatening ecosystem function and human wellbeing. Addressing this crisis effectively and ethically requires transdisciplinary approaches that recognise the complex, culturally embedded interactions between people and freshwater fauna. Existing frameworks for integrating the cultural and ecological dimensions of freshwater fauna are either too general or conceptually misaligned to support practical application, and are often based on binaries such as \u0026lsquo;tangible and intangible\u0026rsquo; that artificially segregate meaning and practice. To bridge this gap, we developed a novel framework for systematically categorising the cultural significance of freshwater fauna, drawing on pre-existing frameworks and human-freshwater-fauna interactions (n\u0026thinsp;=\u0026thinsp;612) from interdisciplinary literature. The resulting framework comprises seven overarching domains \u0026ndash; Consumptive Use; Cultural Landscapes, Sites and Stewardship; Knowledge Systems; Worldviews, Beliefs and Identities; Cultural and Social Practices; Recreation, Leisure and Tourism; and Creative Expression \u0026ndash; each subdivided into defined categories and subcategories. Applicable across diverse cultural, taxonomic and geographic contexts, our framework provides a pragmatic tool to support more holistic and inclusive approaches to freshwater ecosystem research, management, governance, and cross-cultural cooperation.\u003c/p\u003e","manuscriptTitle":"A framework to categorise the cultural significance of freshwater fauna","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-23 12:35:41","doi":"10.21203/rs.3.rs-6830897/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"8ad37114-099f-4ba9-93b1-5d0d413c4ab3","owner":[],"postedDate":"June 23rd, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":50359687,"name":"Scientific community and society/Water resources"},{"id":50359688,"name":"Earth and environmental sciences/Environmental social sciences/Environmental impact"},{"id":50359689,"name":"Scientific community and society/Social sciences/Interdisciplinary studies"}],"tags":[],"updatedAt":"2026-01-12T15:26:51+00:00","versionOfRecord":[],"versionCreatedAt":"2025-06-23 12:35:41","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6830897","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6830897","identity":"rs-6830897","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}