Italian still life paintings as a resource for reconstructing past Mediterranean aquatic biodiversity

We have explored the historical representation of aquatic resources in Italian still-life paintings as an indicator of past aquatic socio-ecosystems. In this study, still-life paintings were used as a selective and powerful lens on the human-nature relationship rather than as a comprehensive archive of aquatic biodiversity. Our focus was on paintings from the Early Modern Era, a period that remains understudied due to the lack of systematic scientific records, particularly for the Mediterranean Sea. Existing historical archives, when available, are often fragmented and country-specific, making access challenging – especially in Italy which only unified in 1861, leaving many historical documents dispersed and largely undigitized.

Using an environmental history approach, we analysed paintings from various Italian regions and periods, examining how artistic representations of aquatic organisms reflect human interactions with aquatic species. Our interpretation of aquatic ecosystems through still-life paintings is shaped by two key filters: a technical filter (fishing techniques, aquaculture and trade) and a socio-cultural filter (culinary preferences, aesthetic choices and symbolism), both of which are developed in the first two sections of our discussion. Taking these filters into account has enabled us to develop a more nuanced ecological interpretation of the variations in the depicted taxa, considering ecological factors such as biogeography, climate and habitat changes, resource overexploitation, and the presence of introduced or cryptic species.

Still-life depictions of aquatic species reflect the exploitation of aquatic resources and technological advances in fishing in Italy. In the Mediterranean Basin, aquatic organisms have long been a central resource for subsistence³⁵. Before the 16^th century, early fisheries in Italy relied heavily on freshwater and anadromous species which were harvested traditionally from artificial ponds and brackish coastal lagoons, especially in the North³⁵. On the other hand, maritime fishing remained a small-scale activity until the 16^th century, primarily aimed at fulfilling the subsistence needs of local communities³⁶. The distinction between fishery activities of inland and coastal localities was clearly visible in the art, with inland paintings associated with freshwater and anadromous species, while marine organisms dominated coastal art. These observations correspond to the most consumed organisms at times when people relied principally on species available in their own locality³⁶. For instance, shellfish were abundantly depicted in paintings from the Adriatic region. Bivalves such as razor pods and Venus clams were hand-harvested traditionally along the North Adriatic coast for centuries³⁷. Similarly, oyster cultivation has long been practiced in the Adriatic, with remains dating back to Roman times³⁸. This custom of farming, together with shellfish consumption, made these taxa prominent subjects in local paintings over the centuries.

The presence of marine organisms, and notably shellfish, was nonetheless extensive in inland paintings throughout the period, highlighting the scale of trade across the Italian peninsula. Trade was an essential part of early fisheries activities, and historical sources indicate prosperous exchanges from inland lakes and rivers towards coastal localities during the 14^th–15^th centuries³⁹. Traditional merchant routes such as the via Aemilia were later followed on a seasonal basis by coastal merchants to transport fresh marine fish to inland localities³⁹. For instance, grey mullets were abundantly represented throughout the studied period in all regions. They were easy to catch in coastal and brackish waters due to their consistent presence and migratory routes⁴⁰, and their high economic value explains the successful trade across regions and throughout centuries⁴¹. In the paintings we observed a noteworthy similarity between inland and Adriatic paintings in the depiction of marine organisms: this similarity reflects regional exchanges which were facilitated by the preservation of these marine organisms which could then more easily be transported across the Po River and along trading routes.

Prior to the 16^th century, fishing in the Mediterranean relied on traditional methods such as the use of hand nets, pots, and traps. These techniques, primarily used for fishing and harvesting freshwater species, coincided with recurrent representations of freshwater organisms in still-life paintings before 1650. During that time, the exploitation of marine resources was dominated by coastal lagoon fishing, and artists often depicted species caught inshore, such as grey mullets. Historical research highlights the prevalence of these local fishing practices in the Mediterranean until the 17^th century, when the economy began to shift toward coastal exploitation³⁹. This shift was driven by new types of fishing gear, such as bottom trawls, longlines, drift nets, and gill nets which greatly increased the exploitation of marine resources⁴². As these methods were adopted, the diversity of catches expanded, which in turn influenced the diversity of species depicted by artists.

Trawling methods targeting benthic species emerged in the 15^th century and had expanded significantly by the 18^th century⁴³. The rise of entire fleets dedicated to bottom trawling led to an increase in both the yield and diversity of catches^44,45, which coincided with a growing depiction of benthic organisms in still-life paintings. This trend was particularly prominent in the Adriatic Sea, where benthic and benthopelagic species were widely represented from the 17^th century onward. The large, shallow continental shelf of the Northern Adriatic facilitated the expansion of trawling, making it common practice by the mid-17^th century³⁹. Artists in the region, inspired by the rich variety found in fishermen’s catches and on fishmongers’ stalls, frequently depicted bottom-dwelling species such as flatfish, stargazers, John Dory (Zeus faber), anglerfish, in addition to benthopelagic sharks and rays. In contrast, the narrow continental shelf along the Liguro-Tyrrhenian coasts of Italy limited the use of trawling, resulting in fewer benthic fish appearing in paintings from this area.

The use of Sardinal drift nets and Tartane trawlers became widespread by the 17^th century in the quest for catching pelagic fish^43,46. However, unlike benthic species, pelagic fish remained underrepresented in Italian still-life paintings, making up overall fewer than 17% of the organisms depicted in art. This trend contrasts with previous observations at the European scale, where painters from Northern Europe increasingly represented pelagic fish across the same studied period²⁶. Two exceptions, however, concern the migratory pelagic tuna and the swordfish. The use of specific types of gear such as tuna madrague to target these large species was widespread by the 17^th century⁴⁷, and is reflected in artistic representations from between 1650 and 1700, suggesting increases in tuna and swordfish catches. However, their depiction sharply declined in the 18^th century, likely reflecting a change in migratory routes towards the open sea^47,48. In Italy in particular, tuna catches significantly dropped from the 17^th century onwards due to climatic change, and using traps ceased to be commercially viable⁴⁹. Conversely, artistic representations of sharks increased over the studied period, likely due to their incidental capture as by-catch in longline fishing. This trend may also correlate with the growing use of commercial longline fishing from the 18^th century onward⁴³.

Socio-cultural factors, including culinary preferences, aesthetic choices, and religious symbolism, also influence the depiction of aquatic resources in art. The representation of aquatic species in still-life paintings attests to the culinary value of these organisms and provides a window onto the tastes of Italian social classes. Most paintings were either commissioned by affluent families and were out of the reach of ordinary people. As a result, the depicted species consisted primarily of fresh, high-quality fish, with few representations of preserved fish (salted, smoked or dried), which lacked the visual appeal of fresh fish but which were more accessible to the lower classes³⁹. Most paintings tended to depict high-value specimens and those appreciated for their culinary interest. For instance, despite their significance in the Mediterranean diet⁴³, small pelagic fishes such as sardines (Sardina pilchardus) and anchovies (Engraulis sp.) appeared in fewer than 10% of the studied paintings. These species were considered low-quality and were sold cheaply to the lower social classes, and were sometimes even deemed unhealthy according to the medical theory of humors³⁹. Similarly, European eels (Anguilla anguilla), though heavily fished from the Medieval period onward⁵⁰, showed a decline in representation through the centuries, linked over time to their association with lower-class consumption⁵¹. Conversely, crabs (Brachyura), mantis shrimp (Squilla mantis), and lobsters (Homarus gammarus) were commonly consumed by all social classes³⁷ and figure frequently in the corpus, especially in the paintings of the Adriatic region. These edible shellfish in particular were artistically displayed in restaurant windows in Venice, as is still the case nowadays. The nutritional and economical significance of the depicted organisms was not analysed in this study but could be proposed for future research.

As underlined by Tribot et al.²⁶, aesthetic choices may also determine the species compositions of still-life paintings. Artists of the Early Modern Period appear to have chosen species based on aesthetic qualities such as colour and shape, without regard for their culinary value. For example, the Recco family of painters frequently included vibrantly coloured species such as gurnard, red mullet, scorpionfish, and seabream (Pagellus sp.), using bold reds to contrast with darker elements in their compositions. Since Antiquity, red has been linked to power and the sacred, and Italian painters, by saturating their works with red, highlighted the preciousness of the depicted marine animals. This emphasis on colour is exemplified in Giovanni Battista Recco’s “Still-life with fish and oysters” (1653) (Nationalmuseum Sweden), where bright colours enhance the visual impact of the painting. In our still-life corpus, the presence of red coloured species was observed in about 65% of paintings, and in up to 79% of paintings originating from Naples, although we were not able to quantify precisely the aesthetic interest for painters. In addition to colour, some species may have been depicted because of their unique and intriguing shapes. Examples include seahorses (Hippocampus guttulatus), flyingfish (Cheilopogon heterurus), sunfish (Mola mola), the red-spotted box crab (Calappa sp.), spiny starfish (Marthasterias glacialis), and the angelshark (Squatina squatina). These species added an element of curiosity and novelty to the paintings, showcasing the artist’s creative choices rather than reflecting common culinary practices.

Still-life paintings from the Early Modern Period also often incorporated religious themes, reflecting the cultural and symbolic significance of certain objects. One notable example is the use of red coral, a material valued in Mediterranean cultures for its alleged protective properties⁵². In 17^th century Italy, “coralline” boats actively fished for coral along the peninsula and around the surrounding islands⁵³. However, by the late 18^th century, declining commercial demand had led to a regulated suspension of coral fishing, which further reduced harvests⁵³. This shift is mirrored in art, particularly in paintings from the Tyrrhenian coast, where red coral appears in compositions featuring mythological figures, emphasizing its symbolic resonance.

Once technical and socio-cultural influences are considered, still-life paintings provide valuable ecological insights by reflecting geographic distribution of species, the effects of climate change and habitat modifications, the overexploitation of emblematic and endangered species, and the presence of introduced and cryptic species. Italian painters displayed a clear propensity for representing locally accessible taxa, which confirms a convergence between the origin of the paintings and the species’ biogeographic area, as demonstrated by Tribot et al.²⁶. In freshwater environments, fisheries essentially targeted abundant nearshore benthopelagic fishes, such as pike, Eurasian perch (Perca fluviatilis), tench and common carp³⁶, all widely depicted in paintings from inland localities, along with other commercially important freshwater taxa. These freshwater taxa were also abundantly represented in Flemish still-life works²⁶, and Italian painters may have been influenced by these representations. Although the historical exploitation of these fishes coincides with still-life representations³⁶, it is not possible to quantify the influence of cultural diffusion of paintings with the dataset at hand. The representation of anadromous taxa, such as sturgeon and trout, was also frequent in paintings originating from both the North of Italy and from the Adriatic coast, in keeping with fishery reports which described these species as abundant in the Po and Tiber rivers and in the upper Adriatic Sea until the 19^th century, but rare in the Liguro-Tyrrhenian Sea⁵⁴.

Regional variations were also notable between the Adriatic and Liguro-Tyrrhenian coasts. Adriatic painters frequently included taxa from sandy habitats, such as common sole, flounder, mantis shrimp, pod razor and the Venus clam in their paintings, mirroring the dominance of sandy shores in the Northern Adriatic. For instance, when compared with Northern European representations²⁶, the mantis shrimp was portrayed almost exclusively in Mediterranean paintings, and notably in the Adriatic, which corresponds to its natural biogeographical distribution. Liguro-Tyrrhenian paintings depicted a great diversity of taxa, including marine molluscs, crustaceans, corals, sea urchins and lampreys. This high diversity was likely due to the variety of habitats found on the Liguro-Tyrrhenian coast, such as sandy banks, Posidonia meadows and rocky habitats. According to historical sources, the variety and abundance of fishes, crustaceans, molluscs, cephalopods and sea urchins was nowhere higher than in fish markets in Naples (Central Tyrrhenian Sea), where they were artistically displayed on stalls^37,40. Liguro-Tyrrhenian artists faithfully represented this diversity with scorpionfish, spiny lobster, seabream, squid, sea urchins, and thorny oysters (Spondylus gaederopus) figuring amongst the most represented taxa.

Climatic changes, particularly during the Little Ice Age, affected the geographic distribution and population dynamics of aquatic organisms. The cold, wet winters prior to the 16^th century were favourable for many freshwater species⁵⁵, in particular for poikilotherm fishes. However, the warming trend after the 17^th century led to population declines. This tendency was observed in our study with the decrease of most freshwater species over time, corresponding to a continent-wide decline, as depicted in a recent study on still-life representations of aquatic organisms across Europe²⁶. This decline was exacerbated by habitat changes due to human intervention. During the studied period and especially around the mid-16^th century, important habitat modifications occurred in inland lakes, rivers and coastal ponds of Europe, leading to the decline of many freshwater fish and invertebrate populations^56,57. By the second half of the 17^th century, drainage of wetlands for the benefit of agriculture led to massive habitat loss, especially in Tuscany, Veneto and Rome³⁹. For instance, the Italian population of sturgeon, a cold-adapted migratory species, began to fall in the 16^th century due to climatic changes and temperature rise, and then endured a further decline by the beginning of the 19^th century⁵⁴ due to overfishing, habitat modification, and pollution. This trend is reflected after 1650 in a corresponding decrease in artistic representations of freshwater and anadromous taxa, such as sturgeon.

Many of the taxa identified in the corpus were commercially exploited for centuries along the Mediterranean coast. Today, several of these species are listed as vulnerable or endangered, and fisheries management plans have been established in recent decades to recover past populations⁵⁸. However, there is a significant gap in scientific records pertaining to their ecological status during the 15^th–18^th centuries. Indeed, most studies that have assessed the ecological condition of vulnerable species are time-based and cover either a distant past (i.e. are based on archaeological remains^8,59) or the relatively recent centuries (i.e. are based on biological sources⁴⁷ or fishery landings⁴⁸). The presence and relative abundance of these species in still-life paintings provides insights into the timeline corresponding to the decline of these vulnerable species, since by the end of the studied period, painters were depicting many fewer of the taxa currently listed as critically endangered, endangered or vulnerable. An example of this is the portrayal of emblematic Mediterranean species such as marine turtles, noble pen shells and red coral, which all declined sharply in paintings made after 1650. Marine turtles have historically been exploited by fisheries for food and trade⁶⁰, which is confirmed by their constant presence in still-life paintings throughout the period studied. They were, however, depicted more frequently between 1650–1700, likely due to the diversification of fishing methods and increased bycatch. The decrease in their representation towards the end of the 18^th century might reflect an early population decline, as was documented a century later in fishing records⁶⁰. Similarly, red coral and the noble pen shell have been exploited since Antiquity in the Mediterranean Sea^52,61. Both species were harvested particularly actively around the Italian peninsula and islands, notably in the south of Italy, culminating in a peak in exploitation during the 17^th century^53,62. While the near exclusive presence of both species in the studied paintings from the Liguro-Tyrrhenian coast corresponds with their geographical range, the observed decrease in their representation after the year 1700 appears to reveal a global disinterest for these species and a potential population early decline.

The high level of detail in still-life paintings has allowed us to identify most species, although some cryptic and introduced species presented us with challenges. For example, Italian pikes are currently represented by two known species, the endemic cisalpine pike (Esox cisalpinus) and the northern pike (E. lucius), both of which are present in several Italian lakes and rivers⁶³. The detailed representation of pikes in the corpus allowed species-level identification by experts for about two thirds of specimens, revealing about 80% cisalpine and 20% northern pike. This result indicates that the representation of northern pikes in Italian paintings might reflect a historical introduction of the northern species from outside the country, leading to introgressive hybridizations among Italian pike populations^64,65. Moreover, the overall higher depiction of endemic pikes suggests that they used to be more abundantly available for fishermen than introduced pikes, yet today they are identified as vulnerable, affected by introgressive hybridization and competition with E. lucius⁶⁶.

Conversely, some species such as the Italian barbels (Barbus plebejus, B. fucini and B. tyberinus sensus lato) were particularly difficult to differentiate in the paintings due to their similar morphological characteristics and frequent hybridization⁶⁷. However, none of the allochthonous barbel that currently occur in Italy such as B. barbus, B.cylcolepis and Luciobarbus graellsii, were identified^68,69. These species have generated introgressive hybridizations with endemic barbel species, leading to conservation issues and attesting their recent introductions^70,71,72,73. The three species were identified in the paintings by experts, although some uncertainties remain. Similarly, we could not precisely differentiate between the Italian native crayfish species belonging to the genera Austropotamobius, nor was it possible to establish if some paintings depicted the allochthonous noble crayfish (Astacus astacus), which was historically introduced into Italian aquaculture in the 19^th century⁷⁴. Earlier signs of its introduction were suggested in paintings, but more detailed research or direct close observation of the paintings by taxonomy experts is required.

Despite their historical significance, some commercially important species are scarcely represented in Italian still-life paintings. Groupers, for instance, were widely portrayed in Roman mosaics²⁰ but are almost absent from Early Modern paintings, probably reflecting an early population decline. The torpedo ray (Torpedo torpedo) was frequently represented in ancient Greek and Roman art due to its various ‘medical’ applications⁷⁵, but is similarly underrepresented in still-life paintings. Its declining use for medical purposes in the 4^th century likely explains this sporadic presence in Early Modern paintings. Similarly, Italian artists depicted very few marine mammals, unlike what can be found in Northern European paintings²⁶. Only one walrus and two seals were found, all three in Arcimboldo’s Water (1586), which is a composite portrait combining aquatic creatures. However, neither of these species belongs to the Mediterranean fauna, as the portrayed seals lack resemblance with the only known Mediterranean species, the Mediterranean monk seal (Monachus monachus). This latter species, although abundant in ancient times, faced population collapse in the 16^th century due to human exploitation⁷⁶, which may explain its absence in the paintings. Yet monk seals were also rarely depicted in early art, with the few known examples including Palaeolithic engravings from the Cosquer cave (27,000–18,500 BCE)⁷⁷, ancient Greek coins (7–6^th century BCE)⁷⁸ and a recently discovered Greek painted vase (6–5^th century BCE)⁷⁸. The relative lack of more recent artistic representation may stem from the reputation of seals in myths and religious writings, where they were categorised as malignant animals among the sea monsters, a belief that stemmed since the spread of Christianity⁷⁸.

In conclusion, the understanding of historical dynamics of human–nature interactions has proven to be challenging. On the one hand, while scientific monitoring has significantly shaped modern understandings of ecosystems, its scope is usually limited to recent decades, offering a relatively short temporal perspective that often overlooks pre-industrial ecological conditions³. On the other hand, more ancient historical archives are not easily accessible and, when available, quantitative information is only partial or imprecise³. These narrow views can inadvertently contribute to the shifting baseline syndrome described by Pauly (1995)⁷⁹. In response to these limitations, researchers have increasingly turned to integrative and interdisciplinary approaches to reconstruct past environments and unravel long-term socio-ecological dynamics^1,2,3. Building on an environmental history approach, our research emphasizes the value of art as a complementary source of ecological and historical data to explore long-term socio-ecological dynamics, especially for periods and places where scientific records are sparse or fragmented. In particular, still-life paintings from the Early Modern Period, renowned for their detailed representations of natural objects, serve as valuable visual archives for examining historical biodiversity. This study offers an analytical framework for interpreting these artworks as source of insight into past socio-economic dynamics, cultural practices, species distributions and ecological changes influenced by climate variability and human activity. At a time when aquatic ecosystems face unprecedented pressures from overexploitation, pollution and climate change, integrating artistic representations with scientific evidence provides a novel and interdisciplinary approach for enriching biodiversity research. By drawing from these diverse sources, we can develop more comprehensive conservation strategies rooted in long-term ecological awareness. Meanwhile, art offers a powerful medium for raising awareness about the urgent need to protect at-risk ecosystems, offering new pathways for engagement in biodiversity conservation.