Anne-Sophie Tribot

Daniel Faget

Thomas Richard

T Changeux

The role of pre-19th century art in conservation biology: An untapped

Keywords: nature connectedness, human-nature interactions, art, mediation tools, aquatic biodiversity Word count: 8151

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The need to integrate nature connectedness into conservation biology

In recent decades, nature connectedness (NC) has been recognized as a key factor in addressing and meeting sustainability challenges [START_REF] Ives | Reconnecting with nature for sustainability[END_REF]. As one of the main levers for the adoption of pro-environmental behaviors (PEBs), NC has been shown to be a relevant tool for promoting the conservation of biodiversity [START_REF] Richardson | The green care code: How nature connectedness and simple activities help explain pro nature conservation behaviours[END_REF]). An increasing number of studies suggest that NC contributes significantly to the adoption of PEBs, i.e. engaging in acts that benefit the natural environment and avoiding acts that harm it [START_REF] Whitburn | Meta analysis of human connection to nature and proenvironmental behavior[END_REF]. In addition, NC actively participates in human happiness and well-being [START_REF] Pritchard | The Relationship Between Nature Connectedness and Eudaimonic Well-Being: A Meta-analysis[END_REF], and therefore concerns both ecology and human health, in line with the sustainable development goals of the United Nations. Nature connectedness can be defined as the individual's subjective feeling of a deep and continuous awareness of the link between self and the rest of nature [START_REF] Zylstra | Connectedness as a Core Conservation Concern: An Interdisciplinary Review of Theory and a Call for Practice[END_REF], and is based on multiple cognitive, sensorial, emotional and experiential dimensions [START_REF] Ives | Reconnecting with nature for sustainability[END_REF]. The evidence linking NC and PEB is mainly based on psychological theories such as the Norm-Activation-Model [START_REF] Schwartz | Normative Influences on Altruism[END_REF], the New Environmental Paradigm [START_REF] Dunlap | The "New Environmental Paradigm[END_REF], the Theory of Planned Behavior [START_REF] Ajzen | L'esthétique du laid, de Hegel à Rosenkranz[END_REF], the Biophilia Hypothesis (Kellert and Wilson, 1993) and the Value-Belief-Norm Theory [START_REF] Stern | A Value-Belief-Norm Theory of Support for Social Movements: The Case of Environmentalism[END_REF]. All these theories highlight that PEBs are sequentially influenced by worldviews, attitudes, values, beliefs and personal norms [START_REF] Oreg | Predicting Proenvironmental Behavior Cross-Nationally: Values, the Theory of Planned Behavior, and Value-Belief-Norm Theory[END_REF][START_REF] Dunlap | The New Environmental Paradigm Scale: From Marginality to Worldwide Use[END_REF][START_REF] Onwezen | The Norm Activation Model: An exploration of the functions of anticipated pride and guilt in pro-environmental behaviour[END_REF][START_REF] De Leeuw | Using the theory of planned behavior to identify key beliefs underlying pro-environmental behavior in high-school students: Implications for educational interventions[END_REF]. Following these theoretical baselines, much research has been dedicated to identifying how the combination of these various factors are associated to PEBs [START_REF] Li | What influences an individual's proenvironmental behavior? A literature review[END_REF] and to NC [START_REF] Pereira | The relationship between connectedness to nature, environmental values, and pro-environmental behaviours[END_REF][START_REF] Whitburn | Meta analysis of human connection to nature and proenvironmental behavior[END_REF]. NC and PEBs are interdisciplinary and multidimensional, so multiple self-assessment scales have been devised for their measurement [START_REF] Hatty | The CN-12: A Brief, Multidimensional Connection With Nature Instrument[END_REF], based on different factors defined according to the psychological theory chosen by the authors [START_REF] Barragan Jason | Human-nature connectedness as a pathway to sustainability: A global meta analysis[END_REF]. Measuring NC and PEBs presents certain limitations: these factors are generally considered socially desirable, and participants may thus overestimate the extent to which they perform these behaviors, or the degree to which they feel connected to nature (de [START_REF] De Leeuw | Using the theory of planned behavior to identify key beliefs underlying pro-environmental behavior in high-school students: Implications for educational interventions[END_REF]. Moreover, if psychometric tools do allow to highlight direct correlations between NC and PEB, they do not inform causality links [START_REF] Whitburn | Meta analysis of human connection to nature and proenvironmental behavior[END_REF]. The development of accurate observational tools and quantitative approaches would constitute a considerable advance in this field [START_REF] Richardson | The green care code: How nature connectedness and simple activities help explain pro nature conservation behaviours[END_REF]. In this perspective, the outcomes of previous studies linking NC and PEBs deserve to be operationally integrated and applied to the field of sustainability science, particularly in conservation biology. To make these generalist tools efficient and informative, they must be tested and adapted to specific contexts and ecosystems for which experts acknowledge the particular socio-ecological issues that need to be taken into account. Moreover, the definition of PEB deserves to be reconsidered in the light of ecological objectives as defined by conservation biology, beyond the prism of environmental psychology. The need to examine pro-nature conservation behaviors (PCBs) more precisely has recently been addressed, as prior research mainly focused on behaviors relating to climate change and resource use, at the expense of behaviors that directly support conservation of habitats and biodiversity (Barbett et al., 2020;[START_REF] Richardson | The green care code: How nature connectedness and simple activities help explain pro nature conservation behaviours[END_REF][START_REF] Barrows | Nature Connectedness, Nonattachment, and Engagement with Nature's Beauty Predict Pro-Nature Conservation Behavior[END_REF]. It thus seems necessary to directly involve conservation biology both in response to this need and in the development of psychometric tools dedicated to PCBs. In this context, this perspective paper aims to incite researchers in conservation biology to take part in projects carried out at the crossroads of arts and sciences. We first develop the ways in which aesthetic experience and the beauty of nature contribute to nature connectedness. After defining nature aesthetic experience across different disciplines, we then discuss the need to integrate conservation biology into interdisciplinary research that examines the link between nature aesthetic experience and connectedness. In this context, we present Western art (in particular environmental art and Art-Sciences) as a means to promote nature connectedness and conservation. Finally, we draw attention to the lack of research exploring European historical art (pre-19 th c.) in this perspective, and demonstrate through different examples the interest these artworks represent for interdisciplinary projects linking art, biological sciences, environmental education and nature connectedness.

2. Engaging with nature through aesthetic experience 2.1 Nature connectedness and engagement with the beauty of nature Facing the current biodiversity crisis and the need to engage people with nature, research in environmental psychology has examined ways to modulate NC. A number of studies have shown that activities involving physical, emotional and sensory engagement with nature seem to have more effect on NC than do knowledge and theoretical education [START_REF] Zylstra | Connectedness as a Core Conservation Concern: An Interdisciplinary Review of Theory and a Call for Practice[END_REF][START_REF] Lumber | Beyond knowing nature: Contact, emotion, compassion, meaning, and beauty are pathways to nature connection[END_REF][START_REF] Ives | Reconnecting with nature for sustainability[END_REF][START_REF] Pritchard | The Relationship Between Nature Connectedness and Eudaimonic Well-Being: A Meta-analysis[END_REF][START_REF] Barragan Jason | Human-nature connectedness as a pathway to sustainability: A global meta analysis[END_REF]. In this perspective, aesthetic and experiential approaches are promising tools to improve NC through nature experiences. The beauty of nature is a fundamental aspect of the human relationship with the natural world and has been extensively studied under the prism of engagement with nature's beauty, i.e. experiencing positive emotional reactions in response to the perceived beauty of nature [START_REF] Zhang | Engagement with natural beauty moderates the positive relation between connectedness with nature and psychological well-being[END_REF]. This particular aspect of human-nature interactions plays a significant role in well-being, NC, and in the adoption of proenvironmental and pro-nature conservation behaviors [START_REF] Zhang | Engagement with natural beauty moderates the positive relation between connectedness with nature and psychological well-being[END_REF][START_REF] Barrows | Nature Connectedness, Nonattachment, and Engagement with Nature's Beauty Predict Pro-Nature Conservation Behavior[END_REF]. In a similar way, a large part of the science of environmental education has been dedicated to the mediation of biodiversity conservation through simple activities that enhance aesthetic experiences and promote engagement with nature's beauty [START_REF] Barrable | Refocusing Environmental Education in the Early Years: A Brief Introduction to a Pedagogy for Connection[END_REF]. Such activities include, for instance, simple exposure to nature [START_REF] Barragan Jason | Human-nature connectedness as a pathway to sustainability: A global meta analysis[END_REF], spending some time each day noticing beautiful things in the nearby natural environment [START_REF] Richardson | The green care code: How nature connectedness and simple activities help explain pro nature conservation behaviours[END_REF], engaging with nature's beauty through artistic creation inspired by nature-based sources [START_REF] Renowden | Exploring integrated ARTSCIENCE experiences to foster nature connectedness through head, heart and hand[END_REF], or viewing visual arts and performances intended to create empathy towards nature and improve knowledge about environmental concerns [START_REF] Curtis | Towards ecological sustainability: observations on the role of the arts[END_REF]. Facing the current general tendency of disconnection with nature [START_REF] Soga | Extinction of experience: the loss of human-nature interactions[END_REF][START_REF] Ives | Reconnecting with nature for sustainability[END_REF], several mediation tools based on indirect exposure have also been examined, revealing a positive effect on NC [START_REF] Barragan Jason | Human-nature connectedness as a pathway to sustainability: A global meta analysis[END_REF]. Regarding the low impact of theoretical education on NC, this could be linked to the method of transmitting scientific knowledge that includes neither affective nor emotional dimensions, two essential facets of NC [START_REF] Barragan Jason | Human-nature connectedness as a pathway to sustainability: A global meta analysis[END_REF].The integration of an approach based on aesthetics and engagement with nature's beauty is thus particularly relevant in environmental education.

Nature and aesthetic experience: a multidisciplinary overview

Nature has long been a topic in aesthetic studies, and defining what is beautiful and pleasurable in nature is a knotty question that cuts across fields as diverse as philosophy, neuroscience, environmental psychology, environmental aesthetics, landscape architecture and human geography. Aesthetics relates to the human ability to judge an object from various perspectives such as emotions, practical use, or experience [START_REF] Shimamura | Aesthetic Science: Connecting Minds, Brains, and Experience[END_REF]. At its origin, Aesthetics was a philosophical field that explored beauty through the study of sensations and emotions involving human perception, particularly by way of Art [START_REF] Brady | Environmental aesthetics: A synthetic review[END_REF]. Beauty is defined in this context as a set of perceptual qualities that generate pleasure or admiration through an aesthetic experience [START_REF] Chatterjee | Neuroaesthetics[END_REF]. However, objects of aesthetic appreciation are not limited to works of art; indeed, aesthetic experiences triggered by nature have long aroused the interest of philosophers [START_REF] Carlson | Environmental aesthetics[END_REF]. Kant, for instance, considered nature to be an object of aesthetic appreciation in itself, and introduced the notions of the Beautiful and the Sublime in nature (Kant, 1790). Beauty is associated with harmony, symmetry and order, and is therefore defined by the intrinsic characteristics of the object [START_REF] Boileau | L'Art poétique[END_REF], while the Sublime arouses admiration and vulnerability, such as may be experienced, for example, when confronting a beautiful landscape beyond the control of the observer [START_REF] Carlson | Aesthetics and the Environment: The Appreciation of Nature, Art and Architecture[END_REF]. During the 18 th century, a third dimension of nature aesthetic judgement was defined: the Picturesque, which connects nature with art to treat nature and landscapes as objects of scenic pleasure [START_REF] Paden | International Association for Environmental Philosophy[END_REF]. This notion greatly inspired 19 th century Dutch painters, especially in their representation of landscapes [START_REF] Paden | International Association for Environmental Philosophy[END_REF]. These philosophical foundations in Aesthetics formed the background for a set of theories in environmental aesthetics beginning in the 20 th century [START_REF] Brady | Environmental aesthetics: A synthetic review[END_REF]. Environmental aesthetics places the aesthetic experience of everyday environments as subject matter, in response to the field of Aesthetics that has long favored art over nature [START_REF] Carlson | Environmental aesthetics[END_REF]. Environmental aesthetics particularly emphasizes the importance of the multisensory immersion of such aesthetic experiences (contrary to the visual and scenic approaches of classical Aesthetics, [START_REF] Berleant | The Aesthetics of Environment[END_REF] and questions whether knowledge is a significant factor of aesthetic valuing [START_REF] Carlson | Aesthetics and the Environment: The Appreciation of Nature, Art and Architecture[END_REF]. Research in philosophical Aesthetics has also shaped the theoretical frameworks of landscape aesthetics and environmental psychology that use an integrative approach in the attempt to identify the biological, psychological, cognitive and socio-cultural origins of the aesthetic experience of nature (see [START_REF] Bourassa | A Paradigm for Landscape Aesthetic[END_REF][START_REF] Kaplan | The experience of Nature, a psychological perspective[END_REF][START_REF] Gobster | The shared landscape: what does aesthetics have to do with ecology?[END_REF]. Regarding the biological and cognitive underpinnings of aesthetic experience, neuro-aesthetics has also provided important insights, even though this field is mainly devoted to the aesthetic judgment of art. [START_REF] Ramachandran | The science of art: A neurological theory of aesthetic experience[END_REF] proposed the 'laws of artistic experience': a list of aesthetic perceptual qualities related to neurophysiological mechanisms. In the same vein, the processing fluency theory of aesthetic pleasure [START_REF] Reber | Processing fluency and aesthetic pleasure: is beauty in the perceiver's processing experience?[END_REF]) integrated these properties into a common framework and linked pleasure to the fluency with which information flows through the cognitive system. As in neuroscience, evolutionary psychology suggests that aesthetic judgement has played a role in human survival [START_REF] Kaplan | The experience of Nature, a psychological perspective[END_REF]. It has been hypothesized that humans have been selected throughout evolutionary history for their extreme proficiency at gathering and processing information from their environment, and that the evaluation of landscape aesthetics is the expression of a selection for simple and reliable information concerning environmental quality [START_REF] Gobster | The shared landscape: what does aesthetics have to do with ecology?[END_REF][START_REF] Kaplan | The experience of Nature, a psychological perspective[END_REF]. Regarding the interdisciplinary literature in aesthetics, a historical dichotomy appears to exist between the aesthetics of art on the one hand and the aesthetics of nature on the other [START_REF] Carlson | Environmental aesthetics[END_REF][START_REF] Barrows | Nature Connectedness, Nonattachment, and Engagement with Nature's Beauty Predict Pro-Nature Conservation Behavior[END_REF]. However, we feel that these approaches should be considered together, because aesthetic experience is a phenomenon that concerns both art and nature. In this sense, integrating the fields of sustainability and biodiversity conservation would constitute a suitable transversal approach for the study of art, nature and aesthetic experience in a more pragmatic and operational way, which could thus be considered as a conservation lever.

The need to integrate nature aesthetics into biodiversity conservation

The interest of integrating the topic of aesthetic experience into conservation biology stems from a simple observation: affective and emotional factors play a fundamental role in the conservation of biodiversity, for we tend to protect and to invest in the conservation of what we love and find beautiful in our environment [START_REF] Gobster | The shared landscape: what does aesthetics have to do with ecology?[END_REF][START_REF] Saunders | Seeing and Doing Conservation Differently: A Discussion of Landscape Aesthetics, Wilderness, and Biodiversity Conservation[END_REF][START_REF] Swaffield | Landscape aesthetic experience and ecosystem services[END_REF]. A telling example dates from the 19 th century, when landscape beauty motivated the creation of the first national parks in the United States: this initiative was greatly influenced by the painters of that time who depicted the wonders of nature [START_REF] Rashkow | Idealizing Inhabited Wilderness: A Revision to the History of Indigenous Peoples and National Parks: Idealizing Inhabited Wilderness[END_REF]. The beauty of nature can therefore be considered as a powerful argument for conservation, provided that the perceived aesthetic value is convergent with the ecological objectives of conservation [START_REF] Gobster | The shared landscape: what does aesthetics have to do with ecology?[END_REF]. Ecosystems with high biodiversity are generally perceived as being the most aesthetic [START_REF] Tribot | Integrating the aesthetic value of landscapes and biological diversity[END_REF]. However, this convergence is not always consistent, in particular regarding human preferences for the conservation of charismatic species [START_REF] Colléony | Human preferences for species conservation: Animal charisma trumps endangered status[END_REF]. Consequently, over the recent decades ecologists have examined how perceived biodiversity affects the aesthetic evaluation of species and ecosystems in an attempt to better understand how the ecology of species influences human interactions with nature and the willingness for conservation [START_REF] Gaston | Personalised Ecology[END_REF]. These assessments are generally based on visual and digital observations of nature (usually photographs) and very few have explored the potential for engaged and engaging nature aesthetic experiences. However, we believe that conservation biology has a role to play when investigating how different facets of nature aesthetic experience affect NC. In this perspective, there is a need to implement into conservation biology the findings of the several disciplines interested in nature aesthetic and NC. In particular, conservation biology would help formalize in this context the notions of 'nature' and biodiversity based on formal metrics, since these concepts remain fuzzy in the human sciences. It is important that the biological sciences take part in research and reflection on the relationships between aesthetic experience, NC, and conservation in order to provide an approach specifically tailored to the conservation of biodiversity. Thus, the development of interdisciplinary research at the crossroads of conservation biology, art and humanities is both urgent and pertinent.

3. Fostering nature connectedness through environmental art Connecting people with nature through aesthetic experience could act as a lever for biodiversity conservation, revealing the need to explore experiences that trigger significant affective and emotional responses. Art is a suitable pathway in this context, because it allows to transcend the cognitive dimension of environmental concerns and also influences people's worldviews and life goals [START_REF] Heras | Realising potentials for arts-based sustainability science[END_REF]. Among recent successful programs connecting art with ecological mediation, many involve environmental art and exhibit a broad range of artistic practices and artworks in any format, including the use of nature as a medium. Since its appearance in the late 1960s, this artistic movement has often been characterized by a political and ethical stance towards nature [START_REF] Brady | Introduction to 'Environmental and Land Art[END_REF][START_REF] Thornes | A Rough Guide to Environmental Art[END_REF] and has aimed overall to raise awareness of environmental issues by fostering an appreciation and interest for nature. Studies in environmental education have examined the potential of such artworks and practices to foster nature connectedness through 'aesthetic learning' (arts-based educational experiences, Heras, 2022). For example, [START_REF] Marks | Environmental art as an innovative medium for environmental education in Biosphere Reserves[END_REF] explored responses to environmental art initiatives in two Biosphere Reserves (Noosa, Australia and North Devon, UK) and revealed that such programs can be used both to enhance understanding of environmental issues and to generate concern. Another way to link art to environmental education, known as Art-Science, involves using scientific data, processes or knowledge as supports for artistic projects [START_REF] Lesen | Science Communication Through Art: Objectives, Challenges, and Outcomes[END_REF]. Aesthetic learning and Art-Science initiatives are often based on participatory practices (see for example [START_REF] Curtis | Towards ecological sustainability: observations on the role of the arts[END_REF][START_REF] Harrower | Species Loss: Exploring Opportunities with Art-Science[END_REF][START_REF] Renowden | Exploring integrated ARTSCIENCE experiences to foster nature connectedness through head, heart and hand[END_REF]. A number of Art-Science artworks intended for large audiences have resulted from collaborations between artists, stakeholders and scientists. One such example is the Cucusonic project, that brought scientists, musicians, farmers, city dwellers and indigenous peoples together to create musical compositions based on bioacoustic recordings in Columbia (see cucusonic.net). Obviously, artistic creation and aesthetic reception do not involve the same amount of physical commitment, but these two facets of art should be considered as complementary and reciprocal [START_REF] Tinio | From artistic creation to aesthetic reception: The mirror model of art[END_REF]. Although participatory approaches allow for a great degree of engagement [START_REF] Lesen | Science Communication Through Art: Objectives, Challenges, and Outcomes[END_REF] they nevertheless have drawbacks: setting up such projects may be more costly in terms of time and resources, and their success may also be highly dependent on the motivation of the participants. Ensuing problems may include sampling bias in the recruitment of participants, or artistic practices that are not always suitable for all participants. For example, [START_REF] Renowden | Exploring integrated ARTSCIENCE experiences to foster nature connectedness through head, heart and hand[END_REF] reported that some participants felt 'infantilized' during a workshop where they were asked to recreate frog songs using everyday objects. Initiatives based on the aesthetic reception of artwork may be experienced by large audiences, where individuals can engage in an aesthetic experience in keeping with their own disposition and sensibilities. Regarding research into art reception (the field of Aesthetics science analyzing the factor structure of aesthetic experiences, [START_REF] Leder | A model of aesthetic appreciation and aesthetic judgments[END_REF][START_REF] Hager | Assessing aesthetic appreciation of visual artworks-The construction of the Art Reception Survey (ARS)[END_REF], few studies have focused on nature aesthetics, as this field mainly concerns classical art [START_REF] Carlson | Environmental aesthetics[END_REF]. Further investigations at the intersection of the psychology of aesthetics, neuro-aesthetics, environmental education and sustainability sciences are therefore required.

4. Beyond environmental art: connecting with nature through pre-19th century art and cultural heritage Art as a tool of mediation for ecological purposes has engaged the interest of artists and researchers since the late 1960s [START_REF] Brady | Introduction to 'Environmental and Land Art[END_REF]. This growing interest in art as a lever for sustainability has mainly focused on post-19 th century art [START_REF] Heras | Realising potentials for arts-based sustainability science[END_REF]. For instance, [START_REF] Lesen | Science Communication Through Art: Objectives, Challenges, and Outcomes[END_REF] reviewed 200 Art-Science projects in the US, and found that all were based on recent artistic projects. Yet focusing only on the post-19 th century period to the detriment of most of our artistic and cultural heritage seems inadequate because these earlier artworks have the potential to elicit a multitude of aesthetic responses and powerful experiences. In addition, pre-19 th c. art provides an opportunity to explore the imagination of our predecessors: examining their art is a way to connect with the intimacy, daily life, and beliefs of their time. In this way, these artworks allow to confront our current relationship to nature in the face of the history of humanity and its rapport with the environment over time. Indeed, ever since humans began producing works of art (50,000 years ago), nature has been a constant source of inspiration through its intrinsic or symbolic value. There is therefore no obvious reason to concentrate research efforts on recent art only: even if a pre-19 th c. artist's direct intention was not to alert to an environmental crisis, their ability to generate a strong aesthetic experience in connection with the representation of nature remains undeniable. Works of art prior to the 19 th c. also constitute a cultural and historical heritage that is worthy of promotion through the prism of environmental education.

A short timeline of European art history until the 19th century

In order to provide relevant perspectives when studying pre-19 th c. art in relation to nature connectedness, nature aesthetics, and conservation biology we will focus on pictorial art, particularly painting, in Western countries. Indeed, these particular artworks (among others) include many representations of nature that have survived throughout time, and they have been thoroughly studied by art historians. To explain our conviction that pre-19 th c. Western art is deserving of more scientific attention, a brief timeline of European pictorial art history may be helpful. Prehistoric art (50,000 BCE to 3,000 BCE) represents the iconographic depiction of humans and animals, whose possible symbolic and linguistic meaning is still under debate. The parietal paintings of this period testify to the fascination and the great interest of humans for the animals that surrounded them. Ancient art (3,000 BC to 6 th c. CE) is defined as the wide variety of artistic and cultural styles developed by ancient societies living during this period. The Romans, for example, represented nature and its wonders with realism, through frescoes and mosaics. Fish and aquatic animals occupy an important place in this iconography, as evidenced by the famous Mosaic of marine life from Pompeii (early 1st century BC). Art of the Medieval period (6 th c. -15 th c.) used symbolism to serve God. The animal was a symbolic figure to which feelings, thoughts, emotions, vices and virtues could be attributed in order to convey the teachings and morality of Christianity (Vézilier-Dussart, 2016). The Early Modern period (15 th c. to late 18 th c.) includes the Renaissance, Mannerism, Baroque art, Rococo and Classicism. The art of this period is mainly figurative. Under the impetus of technical and scientific discoveries, the representation of nature and the understanding of natural phenomena became subject matter. The 18 th c. Enlightenment was marked by the industrial revolution, reflecting a desire to understand the laws of nature in order to better control them. In parallel, the rise of natural theology in the early 18 th c. translated the spectacle of nature into the expression of the Sublime and perfect divine creation, inspiring, for example, an admiration of the seas and oceans which had hitherto aroused fear and aversion since Antiquity [START_REF] Corbin | Le territoire du vide[END_REF]. The Modern period (late 19 th to late 20 th c.), followed by Contemporary art in the 20 th c., is marked by the appearance of photography and Impressionism, leading to the rethinking of the function of art: the goal is no longer to faithfully represent reality, but rather to question and criticize it, and also to reveal reality and its contradictions in a different way. During Romanticism (late 18 th -late 19 th c.) artists depicted pristine wilderness in landscapes charged with emotions to denounce the impact of industrialization and the desire to dominate nature. For example, Caspar David Friedrich's famous Wanderer Above the Sea of Fog (1818) evokes the insignificance of the human being faced with the landscape, blending admiration and incomprehension through an aesthetic of the Sublime. William Turner (1775 -1861) painted landscapes evoking impressive natural phenomena (fire, storms, sunsets or sunrises) inspired by travels, scientific questioning and the industrial revolution. Conversely, Thomas Gainsborough's Mr. and Mrs. Andrews (1750) can be interpreted as an expression of land ownership and capitalist domination over nature and laborers [START_REF] Rose | Feminism and geography: the limits of geographical knowledge[END_REF]. In summary, pre-19 th c. art figuratively depicted the history and lifestyles of civilizations in their diversity, whereas modern and contemporary art take a more reflective and abstract approach with a universal ambition.

Focus on the Early Modern period

A noteworthy example of representations of nature's beauty in pre-19 th c. art is the Early Modern period in Europe (15 th -18 th c.), in which artists represented the visible world with realism and fidelity [START_REF] Gombrich | The story of art[END_REF]. This period is characterized by changes in representations of the world, by religious reforms, great discoveries and by an economic dynamism resulting from numerous innovations. During the Renaissance, arts and sciences operated harmoniously. Driven by their own curiosity, artists and scientists were often one and the same (e.g. Leonardo da Vinci, Albrecht Dürer, Bernard Palissy, etc.), seeking to understand nature and the world from the perspective of the wonders of nature. For this they combined scientific techniques (including the natural sciences) with beautiful and creative representations of their ideas. The discovery of the microscope in the 17 th c. allowed a better understanding of nature and contributed to the appearance of naturalist painters. Among them was Anna Maria Sibylla Merian who traveled to Suriname in 1699 and brought back drawings of plants and insects integrated into their ecosystem, illustrating that a plant is not just an isolated organism, but is part of an ecosystem whose elements interact. During this period, naturalist explorations developed, and artists traveled to report their observations. Travel and discovery thus became recurring themes in 17 th c. artworks. In this way, artists and scientists used exploration and experimentation to further improve a theory, performance or work. The clear separation between the arts (related to creation and passion) and the sciences (associated with rationality) began during the 18 th c. and is still ongoing today [START_REF] Heylighen | Foundations of ArtScience: Formulating the Problem[END_REF]. The Early Modern period is thus of particular interest when considering a reunification of the arts and sciences in the service of environmental education and NC. First, the degree of realism and the observable desire to represent reality permit a relatively reliable biological interpretation of these artworks. Secondly, the living world was treated as a subject in its own right through the emergence of animal representations in painting and the naturalistic approach, offering visions arousing wonder, curiosity, or even terror or pity [START_REF] Vézilier-Dussart | L'odyssée des animaux: les peintres animaliers flamands du XVIIe siècle [exposition, Cassel[END_REF]. This multiplicity of aesthetic responses fits perfectly within the framework of environmental education through the aesthetic experience of nature.

Science based on pre-19 th c. art

Pre-19 th art is generally characterized as being a testimony to past civilizations [START_REF] Gombrich | The story of art[END_REF]; it thus seems logical that scientists today are interested in these artworks in the attempt to better understand our historical connections with nature and the environmental history of living organisms. Researchers have, for example, investigated representations of fruits, flowers, vegetables or cereals in historical art in order to trace evolutionary history regarding selections over time (see for example Benzi and Berliocchi, 1999;[START_REF] De Smet | The Collaboration Between Art History and Genetics -An Unlikely Marriage of Disciplines[END_REF][START_REF] Kazemi-Shahandashti | Ancient Artworks and Crocus Genetics Both Support Saffron's Origin in Early Greece[END_REF]. This kind of initiative, that examines botanical material from art historical sources, requires the close collaboration of biologists and art historians. The former identify the specimens represented; the latter differentiate symbolic and allegorical iconography from reliable representations that were intended to be realistic [START_REF] De Smet | The Collaboration Between Art History and Genetics -An Unlikely Marriage of Disciplines[END_REF]. Other research has shown the interest of historical art in understanding the environmental history of marine mammals (see for example [START_REF] Brito | The wonder whale: a commodity, a monster, a show and an icon[END_REF][START_REF] Mazzoldi | From sea monsters to charismatic megafauna: Changes in perception and use of large marine animals[END_REF] and aquatic biodiversity [START_REF] Guidetti | Ancient art serving marine conservation[END_REF][START_REF] Tribot | Multi-secular and regional trends of aquatic biodiversity in European Early Modern paintings: toward an ecological and historical significance[END_REF]. Finally, researchers have reflected on how painting could inform the European history of climate [START_REF] Metzger | Une géoclimatologie culturelle: Comparaison entre les paysages peints des Hollandais et des Espagnols au « Siècle d'or[END_REF] and air pollution [START_REF] Zerefos | Further evidence of important environmental information content in red-to-green ratios as depicted in paintings by great masters[END_REF]. This research has shown the importance of integrating pre-19 th c. works of art as material in various disciplines in order to improve understanding of variations of the environment and living organisms over time. However, they have not explored their aesthetic reception by contemporaries as a tool for connecting with nature. Here again, we see the need for additional research at the intersection of environmental history and the psychology of aesthetics for the benefit of environmental education. Aesthetic reception could be investigated by estimating quantitatively the visibility of such works in current museums, exhibitions and art reviews, or by conducting quantitative surveys based on psychometric questionnaires dedicated to aesthetic reception.

Connecting with nature inside and outside art museums

In this perspective, museums have a role to play in putting as many people as possible into direct contact with artworks. To achieve this, scientific mediation efforts can be proposed around cross-cutting themes such as biodiversity, allowing the public to apprehend works of art using different approaches. For example, the Anton Dohrn Zoological Station and the Capodimonte Museum of Naples recently proposed an exhibition dedicated to zoology and art history through a collection of Neapolitan Baroque still life paintings of fish and seafood [START_REF] Travaglini | Storia dell'arte e della zoologia marina. Un dialogo a due voci per una cultura accessibile al Museo di Capodimonte[END_REF]. The authors reported that the paintings were effective levers of involvement regarding gastronomy and fishing, themes whose current issues of biodiversity conservation are of particular interest. The authors also stated that the use of a double narrative discourse (natural sciences and art history) as a promotional language reduced the disparities between visitors from different cultural backgrounds, and made the museum more attractive and welcoming for all. This kind of collaboration between scientists and museums is of particular interest in environmental education, since the emotional impact of the art gains strength when paired with a scientific discourse [START_REF] Harrower | Species Loss: Exploring Opportunities with Art-Science[END_REF]. Several other initiatives based on guided tours in museums have been proposed in order to develop engaging learning activities for students by crossing art education and environmental education (see for example [START_REF] Inwood | Creative Approaches to Environmental Learning: Two Perspectives on Teaching Environmental Art Education[END_REF]. However, apart from school programs, museum exhibitions may still remain unattractive or inaccessible to the general public. Faced with this problem, it is thus important that researchers work alongside museums to develop initiatives that bring physically tangible works into general audience mediation events (such as for instance the European Researcher's Night) in order to substitute for in situ aesthetic experiences [START_REF] Inwood | Creative Approaches to Environmental Learning: Two Perspectives on Teaching Environmental Art Education[END_REF]. To this end, some initiatives such as interactive shows and performances (for examples see [START_REF] Lesen | Science Communication Through Art: Objectives, Challenges, and Outcomes[END_REF] can also be exhibited outside the museums.

5. From aesthetic experience to nature connectedness: an interdisciplinary perspective to promote aquatic biodiversity conservation Aquatic ecosystems (oceans, rivers, lakes and wetlands) represent a very pertinent concern in the context of NC through nature aesthetic experience. They aggregate strong human and ecological issues, and as such are included in the 17 Sustainable Development Goals, yet the relationships that humans have with these ecosystems are mostly derived from indirect exposure (typically through food or cultural objects). This may partly explain why perception of these ecosystems gives rise to many beliefs and misconceptions that are often disconnected from ecological realities [START_REF] Langlois | The aesthetic value of reef fishes is globally mismatched to their conservation priorities[END_REF][START_REF] Tribot | Integrating the aesthetic value of landscapes and biological diversity[END_REF]. To remediate this, there is an urgent need to find new ways to connect people with underwater worlds, which are otherwise inaccessible due to remoteness, mobility issues, lack of specific skills such as swimming, or to financial constraints [START_REF] Januchowski-Hartley | Underwater", in: An A to Z of Shadow Places Concepts[END_REF]. With the exception of divers or small-scale fishermen who develop expertise and close relationships with aquatic environments [START_REF] Nightingale | The Embodiment of Nature: Fishing, Emotion, and the Politics of Environmental Values[END_REF], underwater aquatic life does not enter into most people's everyday experiences of nature. The resulting separation between human perception and ecological goals may indeed constitute an obstacle in the acceptance of conservation or restoration programs, and therefore needs to be scientifically addressed. Paradoxically however, these organisms are actually better known and viewed today than they were historically, thanks to technical advances in scuba diving and underwater cameras. We acknowledge that the success of aquariums with the general public [START_REF] Moss | Impact of a global biodiversity education campaign on zoo and aquarium visitors[END_REF], and exhibitions about the deep ocean (e.g., the title Wildlife Photographer of the Year 2021 was awarded to underwater photographer Laurent Ballesta) participate significantly in people's commitment to the conservation of aquatic environments. Pre-19 th c. art is underestimated as an agent for connecting people with the aquatic world: here we propose indirect exposure to these works as a tool for promoting aquatic biodiversity conservation. Art may be seen in this way as an effective means of engaging with these environments and their species by indirect exposure and through aesthetic experience (Fig. 1). While we fully recognize that this aesthetic experience does not replace the true experience of nature, it does, however, enable people to interact and engage with hitherto inaccessible species and environments such as aquatic ecosystems. In the Early Modern period, aquatic species were generally portrayed as bountiful sources of food, diversity and wonder, and were figurations of a boundless might and indomitable nature. Artworks from this period are likely to arouse multiple feelings ranging from curiosity, wonder, or appetite, to pity, disgust or unease [START_REF] Vézilier-Dussart | L'odyssée des animaux: les peintres animaliers flamands du XVIIe siècle [exposition, Cassel[END_REF]. Repulsion or disgust are generally aroused by perceptual characteristics defined as ugliness [START_REF] Monroe | Ugliness[END_REF]. Although ugliness was formalized in Aesthetics during the 19 th c. under the impulse of Hegel [START_REF] Ajzen | L'esthétique du laid, de Hegel à Rosenkranz[END_REF], the Early Modern painters had long depicted ugliness in order to sublimate reality (see for example the figures depicted by Vicenzo Campi (1536 -1591) or Joachim Beuckelaer (1533Beuckelaer ( -1574))). The paintings of Giuseppe Recco (1634 -1695) for example, show kitchens and market stalls with fish and marine organisms that are ugly and perhaps even frightening due to their viscous and rough aspect (see for example The riches of the sea with Neptune, tritons and two nereids, 1684, by Luca Giordano and Giuseppe Recco, Fig. 1). Fishes as subjects were not considered beautiful during the 17 th c., nor were they seen to be particularly interesting. The painters sublimated this a priori ugliness through a high degree of realism and by creating effects of light and texture: obtaining the wet reflections on the fish was a technical and technological feat. Ugliness may also be embellished by originality and curiosity: we take pleasure in seeing curious monstrosities, which are here assembled, ordered, and balanced. Finally, ugliness is sublimated by diversity, which is a principle of beauty [START_REF] Esmein | Beauté et Différenciation[END_REF]: in this way, painters play on the similarities and dissimilarities within the diversity of life. It is noteworthy that shellfish and crustaceans tend to be associated with travel and discovery, and figure as ornaments (see for example Pierre Mignard's The Marquise de Seignelay and two of her sons, 1691). Whether positive or negative, these aesthetic responses elicit pleasure or interest and convey strong emotional, attentional and memorial implications [START_REF] Menninghaus | The Distancing-Embracing model of the enjoyment of negative emotions in art reception[END_REF]. Among the multitude of possible aesthetic responses, observation of these artworks today is thoughtprovoking and helps decipher the evolution of our perception of the environment and its resources over the long term. This increases understanding not only of the linearity of causes and consequences of human impact on nature, but also the changes of human perceptions of nature, and even prompts questions about the variation of human-nature relationships. A simple way to collect the quantitative data necessary for the in-depth examination of reciprocal relations between pre-19 th c. art and NC would be to constitute a database of artworks specifically representing aquatic biodiversity (through participative science programs, such as www.biodivaquart.fr, [START_REF] Tribot | Multi-secular and regional trends of aquatic biodiversity in European Early Modern paintings: toward an ecological and historical significance[END_REF]. A set of analyses of the perceived effect of aquatic NC and PEB on aesthetic responses could then be applied to this corpus, using environmental psychology tools adapted to the specific case of aquatic biodiversity. More broadly, an interdisciplinary study framework based on the links between aesthetic reception [START_REF] Hager | Assessing aesthetic appreciation of visual artworks-The construction of the Art Reception Survey (ARS)[END_REF], environmental psychology (NC and PEB) and ecology (formal identification and biological interpretation of represented organisms) would be particularly informative. To go further, partnerships between research teams and museums would enable to test the effect of aesthetic experiences on future NC and PEBs in relation to aquatic environments through workshops consisting of guided aesthetic paths. At the same time, mediation activities outside museums involving physical media would be beneficial during events dedicated to the general public. Note that although this article has focused on European art as a case study, art from other regions of the world also deserves to be explored through this type of approach, such as the arts of Africa, Oceania or the Americas. For instance, nature and animals occupy a central place in the iconography of the arts and crafts of Pacific Islands (Kjellgren, 2007). Its aesthetic properties are generally expressed as the receptacle of the sacred and immaterial dimensions of the living world, used as both material and source of inspiration, and thus testifying to particular spiritual connections between humans and nature [START_REF] Descola | Les formes du visible: une anthropologie de la figuration[END_REF]. It would be of great interest to carry out cross-cultural surveys on the aesthetic reception of the arts from different regions of the world, in order to learn more about the transversality of the aesthetic experience of nature in different cultural contexts. The development of such interdisciplinary programs represents a potentially rewarding opportunity to bring together scientists from the human and environmental sciences with actors within cultural organizations, while also involving citizens through participatory sciences. We believe this to be of great interest to conservation biology, as it is only by integrating different layers of society into research processes that the goals of improved understanding and concrete application of the links between art, human-nature relationships and behaviors in favor of biodiversity will be achieved.

Conclusions

Experts, managers, policy makers and ecological scientists should fully consider the arts as a viable lever for sustainable development and biodiversity conservation, and thus as a means of initiating changes in belief systems and environmental behaviors. To do this, scientific advances into the relationships between aesthetic experience, nature connectedness and proenvironmental behavior are necessary, and should not be limited to recent and contemporary artworks. We have an opportunity to better understand and quantify the interrelations between aesthetic experience, nature connectedness and behaviors, in particular with regard to environments such as aquatic ecosystems that are less accessible, and for which people's connection depends for the most part on indirect exposure. To conclude this commented review, we believe that these needs can be achieved through the promotion of research and management projects involving very broad disciplines, ranging from social and environmental psychology through the cognitive neurosciences, to art history and the environmental sciences. Finally, we recommend that pre-19 th c. art deserves to be further investigated, because this is an opportunity to better understand historical nature connectedness and to foster stronger relationships with nature. The authors have no competing interests to declare.

Funding sources

This work was supported by Fondation de France; Agence de l'Eau Rhône-Méditerranée-Corse; Région Sud PACA and Institut Méditerranéen pour la Transition Environnementale (Aix-Marseille University). Funding sources were not involved in the preparation of the article.

Figure 1: Art constitutes a relevant lever for enhancing nature connectedness through nature aesthetic experience. However, further examination of the effect of aesthetic experience as derived from indirect exposure to pre-19 th c. art is necessary for the improved understanding of nature connectedness and pro-environmental behaviors. Solid arrows illustrate relationships that have been well established in the literature: 1 = [START_REF] Barrows | Nature Connectedness, Nonattachment, and Engagement with Nature's Beauty Predict Pro-Nature Conservation Behavior[END_REF][START_REF] Lumber | Beyond knowing nature: Contact, emotion, compassion, meaning, and beauty are pathways to nature connection[END_REF]2 = Barrows et al., 2022;[START_REF] Curtis | Towards ecological sustainability: observations on the role of the arts[END_REF][START_REF] Lesen | Science Communication Through Art: Objectives, Challenges, and Outcomes[END_REF]3 = Barbett et al., 2020;[START_REF] Richardson | The green care code: How nature connectedness and simple activities help explain pro nature conservation behaviours[END_REF][START_REF] Whitburn | Meta analysis of human connection to nature and proenvironmental behavior[END_REF]. Dotted arrows represent hypothesized relationships requiring further investigation: the effect of such paintings on NC, PEB and PCB could for instance be examined through questionnaires using the Art Reception Survey (e.g., [START_REF] Hager | Assessing aesthetic appreciation of visual artworks-The construction of the Art Reception Survey (ARS)[END_REF], NC scales (e.g., [START_REF] Hatty | The CN-12: A Brief, Multidimensional Connection With Nature Instrument[END_REF] and PEB/PCB scales (e.g., [START_REF] Barbett | Measuring Actions for Nature: Development and Validation of a Pro-Nature Conservation Behaviour Scale[END_REF]. Through a quantitative approach, statistical relationships between these factors could be modeled. Painting: Luca Giordano, Giuseppe Recco, 1684, The riches of the sea with Neptune, tritons and two nereids, Naples, oil on canvas, 296 x 234 cm, Art Gallery of South Australia.

Acknowledgements

Language assistance and proofreading of this article was provided by Rachel Mackie. We thank the two anonymous reviewers for their comments and recommendations. We also thank Magali Théron, Céline Ventura Teixeira and Pierre Pinchon from Aix-Marseille University for their art history insights.

Declaration of competing interests