The identification of animal species in ancient art is a complex interdisciplinary field that requires collaboration between archaeology, zoology, ecology, art history, anthropology and natural sciences (Binnberg 2017; Evans et al. 2020; Ferronato & Georges 2023). Accurate identifications rely on such interdisciplinarity, as they involve not only the recognition of morphological characteristics but also the interpretation of the animals’ relationship with human societies that depicted or described them (Lonsdale 1979; Werness 2006; Ferronato & Georges 2023; Sanchez et al. 2024). Ancient zoomorphic objects and their depictions provide valuable insights into how ancient peoples perceived local and transregional fauna, the role of animals in daily life and their broader cultural significance (Tidemann & Gosler 2010; Kostuch 2017; Ferronato & Georges 2023). The prominence of animals in antiquity is evident through their widespread appearance in literature and and art, where they served not only as literal depictions but often as symbols connected to religious, mythological or practical uses (Whitley 2001; Robinson 2002; Hurwit 2006; Neer 2012; García 2014; Sagiv 2018; Andreozzi 2020; Dosoo & Galoppin 2022; Faeza & Zahra 2024). Studying these representations sheds light on ancient human understanding of biodiversity and the evolution of human-animal cohabitation in the same ecosystem.

From a zoological perspective, animals depicted in ancient art can be systematically categorized into taxonomic groups based on distinct morphological and anatomical features characteristic of each order or family (Veth et al. 2018; Andreozzi 2020; Evans et al. 2020). However, species-level identification remains challenging and is not widely encountered. Despite this, there are several well-documented cases of successful species identification and classification through ancient iconographic and literary evidence (Böhme & De Haan 2002; Voultsiadou & Tatolas 2005; Johansson 2012; Trinquier 2012; Kitchell 2014; Binnberg 2017; Sidiropoulos et al. 2017; Pareja et al. 2020; Urbani & Youlatos 2020; Böhme & Koppetsch 2021; Guasparri 2022; Sanchez et al. 2024). In Greece, despite its rich ancient culture, there are few contemporary studies employing zooarchaeological data to investigate faunal representation in art, worship or social relations (Bodson 1981; Alexandridou 2022; Mylona 2022).

Among all animal groups, reptiles, particularly snakes, occupy a special place in the history of ancient civilizations, since in many societies they were considered animals with powerful symbolism and were important in ancient mystery cults (Morgan 2008; Håland 2011; Kitchell 2014; Pieraccini 2016). Their depictions range from ancient Egypt to Greek mythology and Eastern traditions to the New World, reflecting their cultural significance as seen in various artifacts (Egli 1982; Walker 2010; Meier & Vidale 2013; Tabarev 2017; Rodríguez Pérez 2020). However, due to the underrepresentation of zoological approaches and the limited knowledge about ancient fauna despite abundant preserved material, understanding how ancient people perceived and interpreted snakes remains challenging (Kostuch 2024).

While depictions of other reptiles and amphibians are generally underrepresented in ancient classical culture, compared to other taxonomic groups (MacKinnon 2014), snakes occupy a prominent place in art and mythology, constituting one of the most recurring and symbolically rich motifs in both art and literary contexts (Bodson & Orr 2002; Salapata 2006; Hawtree 2014; Kitchell 2014). They appear in a wide range of artistic forms, including reliefs on funerary stelae, sculptures, pottery, frescoes, amulets, seals, figurines and decorative elements on various objects used in everyday life or ritual practices. The presence of snakes in ancient Greek and Latin traditions goes beyond decorative or narrative function, they are symbols deeply embedded in the religious and cultural fabric of ancient societies. Their frequent and almost obsessive representation in artistic and mythological scenes attests to the respect and awe they inspired, acting as conveyors of many symbolic associations, as wisdom, rebirth, transformation, afterlife, preservation of beauty, healing, fertility, death, protection, as well as danger, evil or mystery (Sinha 1999; Håland 2011; Philippa-Touchais 2014; Boze 2016; Pieraccini 2016; Rodríguez Pérez 2020; Tsatsou 2024). These attributes resonate not only within the Greek world but also across the broader Mediterranean and global cultural spheres, as the symbolism of the snake demonstrates remarkable diachronic persistence and diffusion, shifting in meaning according to the religious and philosophical worldviews of each era (Håland 2011; Lenders & Janssen 2014; Tabarev 2017; Wright-Carr 2018; Tzirakis 2019). Particularly in early cultic practices and mystery rites, snakes were elevated to sacred beings, often portrayed with supernatural characteristics and interpreted as intermediaries between humans and the divine (Robertson 1983; Morenz 2004; Stothers 2004; Ogden 2014; Pieraccini 2016; Tzirakis 2019).

Despite their clear cultural and religious significance, the scholarly study of snakes in ancient Greek art and literature remains limited, as detailed taxonomic, iconographic analyses identifying the species depicted or referred respectively, are rarely undertaken (Bodson 1981; Kitchell 2014; Böhme & Koppetsch 2021; Danelis 2025). This lack of systematic investigation reveals a significant research gap, which calls for an interdisciplinary approach, combining archaeology, herpetology and art history, to achieve a more comprehensive understanding of the identity and role of reptiles in antiquity.

Ancient Eretria, situated on the western coast of Euboea opposite Attica (Fig. 1), was one of the most prominent city-states of antiquity, with sustained political, military, commercial and cultural activity dating back to the Geometric and Archaic periods (Boardman 1980: 65, 177; Mazarakis Ainian 1987). In the context of the Persian Wars, Eretria allied with Athens and participated in the Ionian Revolt against the Persian Empire. As retribution, the Persians destroyed the city in 490 BC, shortly before the Battle of Marathon (McGregor 1987: 16; Martinez Vazquez 2019). Following its destruction, the city was rebuilt with Athenian support and gradually regained a portion of its former prosperity (Francis & Vickers 1991).

During the Peloponnesian War, Eretria often adopted a pro-Spartan stance, striving to assert its independence from Athenian hegemony—an objective it achieved in 411 BC following a successful revolt (Richardson 1891; Reber 2007). Following this development, Eretria remained autonomous, its urban architecture began to evolve and the city maintained diplomatic relations with Macedonia (Cawkwell 1978). Eretria possessed democratic institutions and a vibrant cultural life (Walker 2004: 228; Verdan 2012). The city’s architectural and urban organization included major public buildings, such as the large theater, the Temples of Apollo, Artemis and Dionysos, necropolises and luxurious private residences adorned with mosaics (Richardson 1891; Ducrey & Metzger 1979a; Reber 1998, 2007; Verdan 2012). Ancient Eretria stands as a vivid example of a regional power with international ties and cultural prestige (Richardson 1891, 1896; Heermance 1896; Ducrey & Metzger 1979a; Mazarakis Ainian 1987; Reber 2007; Ackermann et al. 2021).

During the 1970s, archaeological fieldwork in Eretria, carried out by the Swiss School of Archaeology in Greece, brought to light significant remains of the ancient city. Pierre Ducrey and Christiane Dunant led excavations of several luxurious residences. Among all of them, the House of the Mosaics may not be the largest, but it is certainly the most richly decorated, occupying a prime location, on the lower slopes of the city’s acropolis. This building was constructed in c. 370 BC, likely by a prominent individual, reflecting the cultural and economic prosperity that Eretria experienced during the Late Classical and Hellenistic period, before the house was destroyed by fire in the first third of the third century BC (Ducrey et al. 1993: 48; Reber 2007). The House is an almost perfect square (25 × 26 m) and consists of 15 rooms and courtyards, for private and social use (Ducrey & Metzger 1979a). The most important rooms are Androns 7 and 9; banqueting halls for symposia where the most significant findings were located. In Andron 9, unique mosaics were discovered, giving the house its conventional name, while Andron 7 yielded a wealth of terracotta figurines, including an intricately arranged decorative ensemble (Ducrey & Metzger 1979a, b).

The decoration of Andron 7 features, among other elements, large wall-mounted appliques depicting a Satyr, Pan, a Gorgoneion (Medusa’s Head), three Maenads, a snake and rosettes. Together, these form a cohesive Dionysian-themed composition that was directly affixed to the walls (Ducrey & Metzger 1979b; Mekacher 2003). While the subject matter suggests that the decoration may have been inspired by mystical or religious background, it was more likely an expression of refined aesthetic luxury in line with the tastes of the time (Metzger 1980; Ducrey et al. 1993: 179). Likewise, it functioned as a cultural instrument through which the owner projected an image of himself as both wealthy and sophisticated. The choice of such symbolic motifs during a symposium setting underlines the importance of displaying power, knowledge and spiritual protection (Reber 1998: 117; 2007).

The aim of this study is to identify the snake species depicted in the terracotta snake figurine from Andron 7, through a multidisciplinary approach combining archaeological context, zoological analysis and modern herpetological taxonomy. By establishing a more precise identification, this research contributes to the broader understanding of the representation of the candidate species in ancient art and its symbolic significance.

The taxonomic identification of the animal species represented in the terracotta applique was based on a systematic morphological and ethological analysis of its features, combined with comparative study of known related species occurring locally and nationally (Masseti 2003; Binnberg 2017; Evans et al. 2020; Guasparri 2022). The proposed methodology integrates the description of the archaeological item with the use of scientific identification keys, as well as photographic and illustrated sources from the field of herpetology. A comparable methodological approach has been adopted by Böhme & De Haan (2002) for the identification of a bronze sculpture of a Western Montpellier snake Malpolon ‌monspessulanus ‌(Hermann, 1804), from the Rabat Archaeological Museum (Morocco), originating from the Roman city Volubilis and dating between the first century BC and the third century AD.

The item under study is a terracotta applique (Inv. no. ME 16584) made of clay with white priming layer, representing a limbless reptile in a loosely coiled posture and intertwined multiple times, with morphological features indicating a Squamata ‌Reptile, specifically a snake (Fig. 2). The figurine bears details such as the imprint of scales on the body, as well as a characteristic curvature and posture of the snakes’ body. Its dimensions are 30 × 24.5 cm (Ducrey & Metzger 1979b). Despite the stylized and simplified artistic rendering, which limits the completeness of certain morphological elements, sufficient morphological information is retained for accurate identification of the species. During excavation, the item was found in a fragmented state, with the head and some body parts completely missing (Ducrey & Metzger 1979b). During the conservation phase, only the lost body parts were restored using appropriate clay, while the head was not reconstructed, preserving its documented loss and, consequently, the authenticity of the find.

For our description, following Youlatos et al. (2023), the applique was reexamined at the Archaeological Museum of Eretria, where additional measurements and photographs were taken, in collaboration and under permission of the Ephorate of Antiquities of Euboea, Ministry of Culture, Greece (Fig. 2). The length of the terracotta snake-body is 142 cm with 4 cm diameter. While no cloacal plate was visible, we considered the tapering end of the clay snake-body to be the tail, whose length is 10 cm, with 2.3 cm diameter.

The identification was based on the comparison of the morphological features of the figurine with descriptions of reptile species identification keys, through specialized scientific herpetological manuals (Arnold & Ovenden 2002; Speybroeck et al. 2016) and published authoritative photographs (Valakos et al. 2008; Insacco et al. 2015; Christopoulos et al. 2019, 2022; Christopoulos & Kotselis 2021), focusing mainly on the features that distinguish the families of the suborder Serpentes. Particular attention should be paid to the analysis of posture, shape and body parts such as head and tail, as animal depictions typically bear key characteristics that are distinctive of the species they represent (Bodson 1981).

The main identification criteria included the shape and manner of the curled posture of the body, the morphology and texture of the scales, the shape and size of the tail and the body length to thickness ratios. In addition, the absence of features attributed to other snake species with different morphology was examined. The specific morphological features indicate with high accuracy that the figurine belongs to a species of the snake family Boidae. Furthermore, a comparison with other snake species, which show similar characteristics and a common geographical distribution, was carried out in order to lead to a more reliable identification of the species represented by the figurine. In particular, species whose geographical distribution includes Euboea, the wider Greek territory and adjacent countries were examined.

The absence of a head limits identification, as the head and its scale plates provide the most critical features for differentiation between related species. In addition, stylized elements and damage to the specimen affect the accuracy of conclusions. Nevertheless, distinct morphological and habitus in combination with ethological features and the geographical distribution data are sufficient to support our species identification.

The systematic comparative study of the morphological features of the terracotta figurine led to the identification of the represented species as a member of the family Boidae ‌Gray, 1825, and more specifically of the subfamily Erycinae ‌Bonaparte, 1831. Despite the wear and the loss of the head, the figurine retains sufficient morphological traits to allow for a reliable identification. Furthermore, ethological evidence reinforces our conclusions as to the identification of the species.

One of the most distinctive morphological traits supporting the attribution to Erycinae is the tail structure. The figurine clearly depicts a short, thick tail with a rounded tip, a defining characteristic of Erycinae snakes (Vitt & Caldwell 2013: 609, 610; Speybroeck et al. 2016: 355, 356). This trait strongly differentiates them from other snake families in the same biogeographical zone, in which the tails tend to be thinner, pointed, tapering and often elongated (Vitt & Caldwell 2013: 609, 610). In Erycinae, the tail is so abbreviated and robust that it can even be mistaken for the head, an adaptation linked both to their burrowing habits and to their defensive behaviour intended to deceiving predators (Greene 1973; Baha El Din 2006: 235, 240; O’Shea 2018: 117).

Furthermore, the texture and depiction of the scales, although artistically simplified, are rendered with a smooth texture (no keels) and rounded shape, characteristics typical of the Erycinae subfamily (Tokar & Obst 1993). Although the snake applique should be considered oversized in comparison to natural snake dimensions (which reach generally only up to 80 cm total length), the body proportions, featuring a large diameter relative to length, further support the identification by distinguishing the figurine from thinner snakes, such as those of the Colubridae family (Speybroeck et al. 2016). Additionally, the figurine portrays a robust, muscular body consistent with the stout build characteristic of erycine boa species (Baha El Din 2006). The absence of a head impedes the identification of specific features but does not prevent a broader classification.

Regarding ethological features, such as posture and behaviour, the snake’s body is depicted in a loosely coiled posture, a common motif associated with resting or defensive readiness and one that is frequently observed among members of the Erycinae species. When erycine boa species feel threatened, they coil into a ball and expose their tail as a head mimic (Greene 1973; Vitt & Caldwell 2013: 610). Particularly noteworthy, however, is the inflation of the body, a feature that suggests a defensive behavioural response, most likely intended to deter a potential threat by visually enhancing the animal’s apparent size (Young et al. 2000; specific on Eryx ‌jaculus, ‌Christopoulos et al. 2026).

Comparison with photographic and illustrated modern sources (Arnold & Ovenden 2002: pl. 42; Valakos et al. 2008: 334, 335; Speybroeck et al. 2016: 356; O’Shea 2018: 117) as well as with taxonomically verified specimens in the literature (Anderson 1898: pl. 33, 33.A) and personal archive, confirms that the above morphological features are consistent with snakes of the genus Eryx. Considering the known geographical distribution of Eryx species, the depicted species can be reliably attributed to the Javelin sand boa Eryx ‌jaculus ‌(Linnaeus, 1758) (Fig. 3), which occurs in the area. This species is included in the herpetofauna of Euboea, as well as several other regions across mainland Greece (Valakos et al. 2008: 333-336; Christopoulos et al. 2019, 2022; Pafilis & Maragou 2020: 178, 179; Christopoulos & Kotselis 2021), which further supports the taxonomy of the species. Its depiction in the figurine may reflect iconographic influence derived from established prototypes, or potentially even direct observation of the animal itself in the wild or in captivity by the artist (Thomas 2021), given that ethological elements appear to be represented.

This defensive depiction, combined with the particular posture, gives the figurine a symbolism that may be associated with protective or deterrent roles, probably in the context of cult or magical-religious practices.

The zoological study of archaeological artifacts depicting fauna offers valuable insights into biodiversity and ecological awareness in antiquity. Such representations serve not only as evidence for the presence of specific species in the historical environment but also as indicators of first-hand knowledge about animal morphology and behaviour, as well as their perceived ecological roles, considering that humans lived in a less disturbed natural environment being curious and observing (Bodson 1981; Kitchell 2015).

By examining morphological and behavioural traits, distributional data and the symbolic and artistic conventions of the period, we conclude that the terracotta semi-coiled snake from Andron 7 of the House of the Mosaics in Eretria most likely represents a boid erycine snake, the Javelin sand boa, Eryx ‌jaculus. Admittedly, the absence of the head remains a notable limitation, as species of Eryx exhibit a distinctive cranial morphology that normally provides important diagnostic information. This interdisciplinary framework offers the most reliable path for species identification in ancient representation, helping to avoid common misinterpretations (Binnberg 2017; Binnberg et al. 2021).

Nevertheless, both zoological and archaeological perspectives should be maintained in an open dialogue, enriching our understanding of how ancient people observed, interpreted and artistically represented the natural world. This multiscale approach deepens our understanding of the historical relationship between humans and nature. The scientific analysis of faunal depictions thus extends beyond their archaeological value, informing current debates in conservation biology and environmental history.

– Conceptualization: A.C., T.D.; – data curation: A.C., T.D.;

– formal analysis: A.C., T.D.;

– investigation: A.C., T.D.;

– methodology: A.C., T.D.;

– project administration: T.D.;

– resources: A.C.;

– validation: A.C., T.D.;

– visualization: A.C., T.D.;

– writing – original draft: A.C., T.D.;

– writing – review & editing: A.C., T.D.

The studied finding from the Archaeological Museum of Eretria was examined under the official permission granted for access to the museum by the Ephorate of Antiquities of Euboea, following a relevant application (Incoming Document no. ΥΠΠΟ/266578/11-6-2025). The rights to the depicted monument (Inv. no. ΜΕ 16584) belong to the Greek State and the Ministry of Culture (Law 4858/2021). The monument is under the jurisdiction of the Ephorate of Antiquities of Euboea © Ephorate of Antiquities of Euboea. Hellenic Ministry of Culture. We would like to express our sincere gratitude to the personnel of the Ephorate of Antiquities of Euboea, and in particular to its Head, Dr. Dimitrios N. Christodoulou, Archaeologist, for granting permission to study the finding. We also wish to thank the staff of the Archaeological Museum of Eretria, especially Chief Guard Ms. Stavroula Parissi and Conservator Ms. Fotini Popoli, for facilitating access to, handling of, and the photographic documentation of the finding. Furthermore, we are thankful for the initial guidance regarding administrative information provided by Dr. Alexandra Kostarelli (Archaeologist, Ephorate of Antiquities of Euboea) and Dr. Ioannis Chouliaras, Archaeologist (Head, Ephorate of Antiquities of Thesprotia and Ioannina). Lastly, we would like to thank the three anonymous reviewers for their insightful comments and suggestions on our manuscript.

1. Even popular news networks reproduced this information, e.g. Squires (2015).

2. Similarly, the Dice snake have been proposed to be introduced to Serifos Island, Greece, as resulted from human-mediated translocation, based on molecular analysis of Kornilios & Thanou (2024).

3. In antiquity, narratives that we might now label as mythical were often perceived as factual, which makes it difficult to draw a clear boundary between myth and scientific reality from a modern perspective (Barbara 2012; Kitchell 2015).