Terrestrial Gastropod Traces on Medieval Human Remains: Taphonomic Evidence from Nysa in Western Anatolia

Terrestrial Gastropod Traces on Medieval Human Remains: Taphonomic Evidence from Nysa in Western Anatolia | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Terrestrial Gastropod Traces on Medieval Human Remains: Taphonomic Evidence from Nysa in Western Anatolia Tolga Köroğlu This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7032422/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract This study investigates mollusc-induced taphonomic modifications on human skeletal remains unearthed from 11th–12th century burials at the ancient city of Nysa (Aydın, western Turkey). Taphonomic alterations—such as pitting, linear grooves, and superficial abrasions—were identified on cranial, tibial, rib, and scapular bones. These modifications were evaluated in connection with three terrestrial gastropod species found in situ: Cornu aspersum (= Helix aspersa ), Oxychilus camelinus , and Caracollina lenticula . Macroscopic and stereomicroscopic observations revealed distinct morphological patterns consistent with biological activity, particularly the gnawing and burrowing behavior typical of Cornu aspersum . Endocranial and ectocranial surfaces of cranial bones exhibited small perforations (1–2 mm), while the tibiae showed linear, sequential cavities (2–3 mm), indicating prolonged and directed contact. The spatial association of gastropod shells and modified bones, especially the scapula, strengthens the hypothesis of direct biological interaction. Furthermore, ecological traits and seasonal activity cycles of the gastropods support the inference that the burials may have occurred during spring or autumn, when gastropod activity peaks. This study documents a rarely addressed taphonomic phenomenon and emphasizes the interpretive potential of gastropod-bone interactions for reconstructing burial conditions, microenvironmental dynamics, and seasonality. By integrating malacological, archaeological, and taphonomic evidence, the research contributes to a more nuanced understanding of inhumation processes and highlights the need to consider invertebrate activity as a significant factor in post-mortem bone alteration. Taphonomy Bioarchaeology Nysa Burial Forensic Archaeology Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 1. Introduction Taphonomic analyses facilitate comprehension of the post-mortem physical, chemical and biological processes undergone by skeletal materials obtained during archaeological excavations. As Lyman ( 1994 ) and Behrensmeyer ( 1978 ) have demonstrated, these analyses assess not only the state of preservation of the remains, but also provide information on such factors as burial conditions, the environmental characteristics of the burial environment, and microbiological interactions. In this context, the presence of biological traces on skeletal material, particularly those left by invertebrates, represents a significant source of data for the comprehension of taphonomic dynamics within the burial environment (Fernández-Jalvo & Andrews, 2016 ). The scientific study of the remains of organisms from the past, known as archaeology, is a field referred to as taphonomy. This branch of science is important in helping us to understand the environmental and cultural contexts of human history. It does this by examining how the remains of organisms from the past were preserved, transported and changed. In this context, the role of gastropods is twofold: firstly, they play an important role in the reconstruction of the natural environment, and secondly, they are important in the identification of traces of human activity. Nevertheless, the potential information these remains hold has been overlooked for a considerable period, with interpretations frequently based on superficial observations or incomplete methodologies. A significant challenge encountered during the archaeological evaluation of gastropods pertains to the ambiguity surrounding the provenance of their remains, namely whether they were transported to the site by cultural or natural processes. This distinction forms the basis of taphonomic analyses (Bobrowsky, 1984 ). In his comprehensive review (1984), Bobrowsky provides a detailed account of the place of gastropods in archaeomalacology, their historical development and methodological problems. In particular, four main ways in which they appear in the archaeological record are highlighted: as a food source, through natural populations, by accidental transport, or for cultural purposes (e.g. as ornaments). These findings imply that taphonomic analyses should be considered not only in a physical but also in a cultural context. These findings imply that taphonomic analyses should be considered not only in a physical but also in a cultural context. Furthermore, the preservation potential of terrestrial and freshwater gastropods may vary due to their exposure to different postmortem processes, necessitating the refinement of taphonomic analyses on a species-by-species basis. Although the influence of invertebrates has long played a secondary role in archaeological taphonomy, it has recently been established that molluscs such as terrestrial gastropods can interact with skeletal material and leave morphologically distinguishable traces on the bone surface (Trueman & Martill, 2002 ). These include superficial pitting, irregular abrasions, linear gnaw marks and microscopic dissolution patterns. However, studies that systematically identify and taxonomically classify these types of scars are very limited. This has led to the attribution of these scars to other factors (microbial decomposition, root scars, erosion, etc.). In the field of gastropod taphonomy, Galvao et al. (2015) conducted a study on the evaluation of gastropod traces in forensic medicine. The study commenced with the discovery of a human skeleton buried for a period of five years in Brasília, the capital city of Brazil. A total of 20 snail shells belonging to the Allopeas micra species were found inside the femur and humerus bones. The observation that all of the shells were found to be in the same developmental stage indicates the likelihood that they belonged to the same generation and entered the bone at approximately the same time. This finding suggests that the snails may have been actively digging into the soil and entering the bones, where they would have continued to live but eventually died. The study emphasises that the snails do not directly intervene in the body tissue, but adapt to the ecological changes in the burial environment. The objective of this study is to analyse the molluscan taphonomic traces present on the skeletal material of an individual unearthed during excavations at the ancient city of Nysa in Aydin province, western Turkey. The graves are located within a necropolis that dates to the 11th-12th century, and the soil structure is characterised by its dry, fine-grained texture and relative paucity of organic matter. The gastropods Cornu aspersum (= Helix aspersa ), Oxychilus camelinus and Caracollina lenticula , which were found at the same level as the skeleton of the studied individual, represent both the grave microfauna and are considered as possible perpetrators of the gnawing and superficial burrowing marks observed on the skeleton. In consideration of the ecological behaviour exhibited by these species, the inactivation of Cornu aspersum during both summer and winter months offers an indirect indication that the sepulchre was most likely accessed during the spring or autumn seasons, periods that are characterised by elevated levels of humidity and mild temperatures (Dahirel et al., 2015 ; Yom-Tov, 1971 ). Consequently, this study not only documents a rare taphonomic phenomenon, but also has the potential to comment on the timing of burials and the biota of the burial environment. 2. Materials and Methods The skeletal material examined in this study was recovered from a grave dating to the 11th-12th century during archaeological excavations at the ancient city of Nysa in the province of Aydin in western Turkey. The grave is located in the eastern part of the necropolis area, in a stratigraphic level with a dry and fine-grained soil structure, and the burials are individual and primary. Despite the partial absence of the skeletons of the individuals exhumed from the grave, the majority of the cranial and post-cranial elements are preserved. The taphonomic marks observed on the skeletal material were documented primarily by macroscopic methods, with the naked eye and using a hand magnifier (10×). The presence of linear and irregular grooves, pitting, and superficial abrasions on the bone surface was meticulously catalogued and categorised in accordance with the taphonomic identification system proposed by Fernández-Jalvo and Andrews ( 2016 ). The anatomical regions in which the lesions were found were meticulously documented, and their morphological characteristics were measured and photographed. For micro-morphological evaluation, a number of lesions were analysed under a stereo microscope (40x), with the bone surface structure being observed in detail. In addition, the skeletal remains of three distinct terrestrial gastropod species were documented within the graves at the level at which the skeleton was discovered. The species under consideration are Cornu aspersum (= Helix aspersa ), Oxychilus camelinus and Caracollina lenticula . In the identification of species, local malacological sources and ecological species identifiers were utilised (Yildirim et al., 2006; Schileyko, 2003 ). The biological potential, morphological structure and feeding behaviour of these species, which may form skeletal marks, were evaluated in the light of the literature. The findings were evaluated in comparison with the relevant taphonomic literature and interpreted by distinguishing them from other possible taphonomic factors (root traces, microorganism decomposition, mechanical abrasion). 3. Results The taphonomic scars identified in the study are considered to be of molluscan origin. These were found on various bones belonging to different individuals. The morphology, distribution and size of the marks exhibited significant inter-individual variability. In all specimens, the marks were found to be superficial gnawing, pitting, burrowing and erosion-like. The potential effects of the identified species Cornu aspersum, Oxychilus camelinus and Caracollina lenticula were taken into account in the evaluation of these scars. On the cranium of one individual, taphonomic scars consisting of small pits and punctures with rounded and irregular edges, measuring 1–2 mm in diameter, were found on both the ectocranial and endocranial surfaces. A more comprehensive documentation of these scars was conducted on the inner surfaces of the frontal and parietal bones. The marks are deep enough to penetrate the cortical bone rather than being superficial. Irregular erosion-like abrasions were also observed on the ectocranial surface in the vicinity of some of the scars. These findings indicate long-term biological activity of molluscs in a humid environment. Large and distinct scars were discovered on the right tibia of a different individual, especially in the periosteal region. Their width is approximately 2–3 millimetres, and they are arranged in linear rows along the surface. Each groove manifests a round or oval morphology, meticulously organised such that it is subtly depressed into the bone surface. The sequential arrangement of these tracks is consistent with the regular gnawing of bone by a gastropod while it moves in a specific orientation. Microscopic analyses of the tracks also documented surface softening and absorption. Taphonomic scars of a similar nature, albeit more superficial and irregularly distributed, were observed on the ribs of another individual. The scars are particularly concentrated on the posterior rib segments and manifest as shallow, pitted grooves with a diameter of 1–3 mm. In some cases, the scars were found to be in close proximity to each other, and upon closer inspection, it was observed that these scars had merged to form larger superficial abrasions. These morphological structures suggest that gastropods were concentrated in a specific area and probably had prolonged contact. On the scapula bone of an individual subject, several pits with irregular edges and measuring 2–3 mm in diameter were found on the surface. In the grave from which this bone was recovered, a gastropod shell was documented in situ next to the scapula. The shell is morphologically well preserved, and a taxonomic examination revealed that the grave of this individual belongs to the species Oxychilus camelinus. This provides a robust environmental context in which the traces can be directly associated with gastropod biological activity. Table 1 General Characteristics of the Findings of Tafonomy Formations Bones Thrace Type Size (mm) Surface Interpretation Skull Perforation (pits) 1–2 Endo/Ecto Progressive biological wear Tibia Sequential Cavity 2–3 Periosteal Regular Gastropod Gnawing The Rib Superficial Pit 1–3 Plevnal Irregular Activity Scapula Adherent Shell 2–3 Lateral Surface Spatial relationship with shell 4. Discussion The spatial correlation of the taphonomic traces observed on the human skeletons obtained from the graves in this study with the terrestrial gastropod species found at the grave levels suggests that these traces may be of biological origin. The species under scrutiny, Cornu aspersum (= Helix aspersa ), Oxychilus camelinus and Caracollina lenticula , offer highly significant data within this context with regard to their ecological characteristics. Cornu aspersum is a species that is particularly prevalent in humid and temperate regions, and it is characteristic of the Mediterranean climate. It has been documented to occur naturally on the western and southern coasts of Turkey (Herbert, 2010 ; Yıldırım et al., 2006 ). In addition to feeding on plant material, it has been observed that the species tends to gnaw bony surfaces to meet its calcium needs (Trueman & Martill, 2002 ). It is evident that this species exhibits heightened activity during the spring and autumn seasons, entering periods of estivation and hibernation during summer drought and winter cold (Charrier & Daguzan, 1980 ). This behavioural pattern lends support to the hypothesis that the gnaw marks observed on the tibia and cranium occurred during the active seasons and that individuals were most likely buried in spring or autumn. The regularly arranged burrow marks, especially on the tibia, are consistent with the oriented gnawing behaviour exhibited by this species. Oxychilus camelinus is another species of land snail found in western and southeastern Turkey, and is adapted to moist and shaded areas (Heller et al., 1997 ). The feeding of this species on microbial film and decaying organic matter has been demonstrated to limit its capacity to directly damage the bone surface. However, it may support the bioerosion process occurring around the bone. The irregular and shallow pits, particularly evident on the rib bones, may be attributable to the indirect contribution of this species, a phenomenon that can be attributed to the microhabitat effect. Caracollina lenticula , a smaller species, inhabits rocky and moist areas, especially in the Aegean Region. It has been observed that the organism feeds on organic sediments and layers of microorganisms. This observation indicates the presence of intense humidity in the burial environment. The documentation of this specimen alongside Cornu aspersum bark, particularly in the context of the grave where the scapula bone was unearthed, provides a direct spatial context for the origin of the traces observed on the bone. In consideration of the life cycles of the aforementioned species, which average at between two and five years, the traces observed on the bone surface appear to be the result of protracted biological activity within the inhumation environment. Furthermore, the seasonal behaviour exhibited by the species offers indirect insights into the timing of burial. The observation that Cornu aspersum demonstrates the highest levels of activity during both spring and autumn supports the hypothesis that burials may have occurred during these seasons. The observations made on the endocranial surface indicate that molluscs gained entry to the cranial cavities and established long-term biological contact. The active involvement of these three species in the microecology of the grave provides information on taphonomic aspects, as well as the moisture balance, soil structure and seasonal climatic conditions of the grave environment. The rarity of the documentation of gastropod-derived taphonomic traces on human skeletons in the literature (see Fernández-Jalvo & Andrews, 2016 ) renders this study a significant exemplar within this field. In consideration of the morphology, dietary biology and environmental sensitivity of gastropod species, it is evident that these species may have played an active role in intra-grave biological processes. The taphonomic traces documented in this study are most likely the result of biological activity by molluscs, given their morphological structure, distribution on bone, and spatial relationships with gastropod species found in the grave contents. In particular, the gnawing behaviour of species such as Cornu aspersum on hard surfaces to meet their calcium carbonate needs has been previously observed in both natural and experimental environments (Trueman & Martill, 2002 ; Yom-Tov, 1971 ). The probability of biological interactions in graves increases during the spring and autumn months when this species is active outside the estivation and hibernation periods. The presence of pore and pit marks on both the endocranial and ectocranial surfaces of the skull is a distinctive feature that has not been previously documented in the literature. In particular, in the endocranial regions, these marks indicate that the organisms had a long-term presence in the skeletal cavities and had access to both surfaces of the bone. Furthermore, the sequential burrow marks on the tibia indicate that Cornu aspersum gnawed while moving in a controlled orientation. This regularity lends support to the hypothesis that the tracks are not random, but rather based on biological behaviour. It is important to note the potential for such traces to be misinterpreted as other taphonomic factors, including but not limited to root action, insect activity, or chemical dissolution. However, the presence of a gastropod shell alongside a scapula bone in an in situ grave suggests that this effect may not be accidental, but rather the result of a systematic interaction specific to the grave microenvironment. In a similar vein, Marchiafava et al. (2015) conducted an experimental study that demonstrated the presence of molluscs in graves can leave marks on bone surfaces. However, the fact that the traces detected in this study exhibit a more intense and different distribution in terms of diameter and depth reveals the uniqueness of the Nysa sample. In the existing literature, the taphonomic effects of molluscs are predominantly discussed in the context of animal bones or prehistoric contexts, while their effects on human skeletons are seldom documented (Fernández-Jalvo & Andrews, 2016 ). Consequently, the data presented here are significant for two reasons. Firstly, they serve to document a new group of agents in the field of archaeological taphonomy. Secondly, they facilitate a more profound comprehension of inhumation biological processes. 5. Conclusions This study revealed that the taphonomic marks observed on human skeletons recovered from 11th-12th century tombs of the ancient city of Nysa may be related to the biological activities of terrestrial gastropods. The species Cornu aspersum, Oxychilus camelinus and Caracollina lenticula were identified as potential factors in the formation of these marks, and were evaluated in terms of both their morphological and ecological characteristics. In particular, the coincidence between the feeding behaviour and seasonal activities of the species with the punctures, sequential burrows and superficial erosion-like scars seen on the bone surface indicates that these scars may be of gastropod origin. The presence of traces on various bone regions, including the skull, tibia, ribs, and scapula, has enabled the formulation of inferences concerning not only the biological processes within the grave, but also the period of burial, the environmental moisture balance, and the post-burial microfaunal dynamics. Cornu aspersum has been observed to exhibit heightened activity during the spring and autumn months, thereby supporting the hypothesis that individuals may have been interred during these periods. The fact that the traces left by gastropods on skeletal material in archaeological contexts are often overlooked increases the importance of this study. In this context, the study introduces a novel perspective to research on both human taphonomy and funerary microfauna, thereby drawing attention to the taphonomic role of terrestrial molluscs. In the future, further studies supported by experimental taphonomy, microscopic analyses and environmental data will serve to reinforce the accuracy of these findings and provide more refined results in taphonomic interpretations. 6. Limitations Despite the fact that the present study sets out to establish the relationship between molluscan taphonomic tracks and their possible perpetrators, it is important to consider the limitations of the study. Firstly, the fact that the traces were evaluated solely on the basis of morphological observation and contextual association limits the precise establishment of a cause-effect relationship. In particular, microscopic or chemical trace analyses were not performed in order to determine the gastropod origin of the pits and abrasions seen on the bone surfaces, and compositional characterisation of the traces was not possible. Furthermore, given the potential for other biological agents (e.g. microorganisms, insects, root systems) present in the grave contents to leave similar traces on the bones, the necessity for additional comparative samples is apparent, with the aim of excluding alternative taphonomic factors. In this context, the hypothesis that the present traces have a molluscan origin is regarded as the most probable scenario in view of the extant data; nevertheless, it is acknowledged that alternative explanations cannot be wholly excluded. In subsequent studies, experimental taphonomy research in controlled laboratory settings can be utilised to more systematically document how different gastropod species leave marks on the bone surface. Furthermore, the employment of advanced imaging techniques, such as scanning electron microscopy (SEM) or micro-CT scans, facilitates a more detailed analysis of the micromorphology of the tracks. Furthermore, the impact of environmental factors associated with the burial environment, including soil chemistry, moisture content and seasonal variability, on trace formation will be a pivotal area of focus in future research endeavours. Finally, the under-documentation of such traces in the literature necessitates more careful monitoring of the taphonomic impact of molluscs in archaeological excavations and systematic recording of microfauna data. Declarations Author Contribution All work belongs to the responsible author. Acknowledgements I would like to express my gratitude to Professor Ümit Kebapçı for his assistance with the identification of gastropod species and Professor Serdar Hakan Öztaner, head of the Nysa Excavation, for his support. References Behrensmeyer, A. K. (1978). Taphonomic and Ecologic Information from Bone Weathering. Paleobiology , 4(2), 150-162. Doi:10.1017/S0094837300005820 Bobrowsky, P. T. (1984). The History and Science of Gastropods in Archaeology. American Antiquity , 49(1), 77—93. https://doi.org/10.2307/280513 Charrier, M., & Daguzan, J. (1980). Consommation alimentaire, production et bilan énergétique chez Helix aspersa müller (gastéropode pulmoné terrestre) [Food consumption, production and energy evaluation in Helix aspersa müller (a terrestrial pulmonated gasteropod)]. Annales de la nutrition et de l'alimentation , 34 (1), 147–166. Dahirel, M., Olivier, E., Guiller, A., Martin, M.-C., Madec, L. and Ansart, A. (2015), Movement propensity and ability correlate with ecological specialization in European land snails: comparative analysis of a dispersal syndrome. J Anim Ecol , 84: 228-238. https://doi.org/10.1111/1365-2656.12276 Fernández-Jalvo, Y., & Andrews, P. (2016). Atlas of Taphonomic Identifications. 1001+ Images of Fossil and Recent Mammal Bone Modification. The Netherlands: Springer. https://doi.org/10.1007/978-94-017-7432-1 Galvão, M. F., Pujol-Luz, J. R., de Assis Pujol-Luz, C. V., de Rosa, C. T., Simone, L. R., Báo, S. N., Barros-Cordeiro, K. B., Pessoa, L., & Bissacot, G. (2015). Shells and Bones: A Forensic Medicine Study of the Association of Terrestrial Snail Allopeas micra with Buried Human Remains in Brazil. Journal of forensic sciences , 60 (5), 1369–1372. https://doi.org/10.1111/1556-4029.12882 Heller, J., Sivan, N. and Hodgson, A.N. (1997), Reproductive biology and population dynamics of an ovoviviparous land snail, Lauria cylindracea (Pupillidae). Journal of Zoology , 243: 263-280. https://doi.org/10.1111/j.1469-7998.1997.tb02781.x Herbert, D.G. (2010). The introduced terrestrial mollusca of South Africa. SANBI, Biodiversity Series, 15 Lyman, R. L. (1994). Vertebrate Taphonomy. Cambridge: Cambridge University Press. Schileyko, A. A. (2003). Treatise on Recent terrestrial pulmonate molluscs. Part 10. Ariophantidae, Ostracolethidae, Ryssotidae, Milacidae, Dyakiidae, Staffordiidae, Gastrodontidae, Zonitidae, Daudebardiidae, Parmacellidae. Ruthenica. Supplement 2: 1309-1466. Trueman, C.N. and Martill, D.M. (2002), The long–term survival of bone: the role of bioerosion. Archaeometry, 44: 371-382. https://doi.org/10.1111/1475-4754.t01-1-00070 Yom-Tov, T. (1971). Body Temperature And Lıght Reflectance In Two Desert Snaıls, Journal of Molluscan Studies , Volume 39, Issue 4, 319–326, https://doi.org/10.1093/oxfordjournals.mollus.a065111 Yıldırım, M.Z., Koca S.B., Kebapçı, Ü. (2006). Supplement to the Prosobranchia (Mollusca: Gastropoda) Fauna of Fresh and Brackish Waters of Turkey. Turkish Journal of Zoology . 30: 197-204. (PDF) The gastropod fauna and their abundance, and some physicochemical parameters of Lake Gölbaşı (Hatay, Turkey) . Additional Declarations No competing interests reported. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7032422","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":479959007,"identity":"7fa7b90c-c222-4200-abbf-235ac3958343","order_by":0,"name":"Tolga Köroğlu","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA6ElEQVRIie3RMQrCMBSA4RcCrUOqa6SoV1ALoqh4lhLoVO9QEOISne3kLdyl0MkbZBG8QNyKdDCtg4jQODrkH8Ir9CMkAbDZ/jFSr1Nw3M35qvTouCbxIhTaJGejQ0Xwr6RP44lff5jIimS5Ug8achpF/uJxGrQxIHWPG3bxOEvTvSbkls3XezniGHA3PTWQDgmwJzRxo1CuhUSaONj7iUA89GdCrszEEwEmhSateOxDIUMzueQMpQkNeHXJu0QyjtGm8SyuYBmoctE7bvVTFqVcVoO6N5A6xD8GlBj+ryq/BpvNZrO9ewIGpExnumAomAAAAABJRU5ErkJggg==","orcid":"","institution":"University of Tübingen","correspondingAuthor":true,"prefix":"","firstName":"Tolga","middleName":"","lastName":"Köroğlu","suffix":""}],"badges":[],"createdAt":"2025-07-02 20:23:13","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7032422/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7032422/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":87545916,"identity":"d1a16ba5-ee16-4157-aff2-fee57a273282","added_by":"auto","created_at":"2025-07-25 04:59:15","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":116500,"visible":true,"origin":"","legend":"\u003cp\u003eTaphonomy Marks on the Skull. As illustrated on the left, the ectocranial region exhibits perforations of a specific configuration. It is evident that analogous traces are present on the inner surfaces of the skull of the aforementioned individual.\u003c/p\u003e","description":"","filename":"image1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7032422/v1/f02132e925454f64b67c2ffa.jpg"},{"id":87545917,"identity":"57547c96-15bd-4b28-b3ad-2348bc9eed54","added_by":"auto","created_at":"2025-07-25 04:59:15","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":250322,"visible":true,"origin":"","legend":"\u003cp\u003eLinear Taphonomy Traces Detected on Tibia.\u003c/p\u003e","description":"","filename":"image2.png","url":"https://assets-eu.researchsquare.com/files/rs-7032422/v1/31f3ff971b528795ab20b251.png"},{"id":87546233,"identity":"cdec063c-f888-4e10-9e0a-cb4a48ef41c0","added_by":"auto","created_at":"2025-07-25 05:07:15","extension":"jpeg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":191907,"visible":true,"origin":"","legend":"\u003cp\u003eTraces of Taphonomy in the Perforated Form of the Pleural Face of the Rib.\u003c/p\u003e","description":"","filename":"image3.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7032422/v1/e28269fa8963cdf225b66eed.jpeg"},{"id":87546236,"identity":"2ae3e023-06da-4ec3-b0a1-0f66cf5a5aab","added_by":"auto","created_at":"2025-07-25 05:07:15","extension":"jpeg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":246404,"visible":true,"origin":"","legend":"\u003cp\u003eThe shell of the snail Oxychilus camelinus was found in situ, adhering to the outer surface of the scapula.\u003c/p\u003e","description":"","filename":"image4.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7032422/v1/2ef2b2e8b62652134b4c14db.jpeg"},{"id":88505226,"identity":"43563c0b-8b67-4d8e-a844-f3edfe2665ea","added_by":"auto","created_at":"2025-08-07 07:21:32","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":157052,"visible":true,"origin":"","legend":"\u003cp\u003eShell Finds of Cornu aspersum Species\u003c/p\u003e","description":"","filename":"image5.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7032422/v1/a0b401c0ee85f543b65f988b.jpg"},{"id":87545929,"identity":"70d9fc9d-7862-4baa-915f-94a2757e0ac9","added_by":"auto","created_at":"2025-07-25 04:59:15","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":85258,"visible":true,"origin":"","legend":"\u003cp\u003eShell Finds of Oxychilus camelinus Species\u003c/p\u003e","description":"","filename":"image6.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7032422/v1/c7facff81176feba32d137f4.jpg"},{"id":87545928,"identity":"341aebea-3dca-4303-92c9-652b66a71c5e","added_by":"auto","created_at":"2025-07-25 04:59:15","extension":"jpg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":127648,"visible":true,"origin":"","legend":"\u003cp\u003eShell Finds of Caracollina lenticula Species\u003c/p\u003e","description":"","filename":"image7.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7032422/v1/3ec684fada846ab83fa78be8.jpg"},{"id":95759380,"identity":"bab118d8-5763-4c27-a811-2fa6874ec1a0","added_by":"auto","created_at":"2025-11-12 17:23:41","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1683295,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7032422/v1/3197fa0a-44f3-46f5-8d36-172e990bc61a.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Terrestrial Gastropod Traces on Medieval Human Remains: Taphonomic Evidence from Nysa in Western Anatolia","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eTaphonomic analyses facilitate comprehension of the post-mortem physical, chemical and biological processes undergone by skeletal materials obtained during archaeological excavations. As Lyman (\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1994\u003c/span\u003e) and Behrensmeyer (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1978\u003c/span\u003e) have demonstrated, these analyses assess not only the state of preservation of the remains, but also provide information on such factors as burial conditions, the environmental characteristics of the burial environment, and microbiological interactions. In this context, the presence of biological traces on skeletal material, particularly those left by invertebrates, represents a significant source of data for the comprehension of taphonomic dynamics within the burial environment (Fern\u0026aacute;ndez-Jalvo \u0026amp; Andrews, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe scientific study of the remains of organisms from the past, known as archaeology, is a field referred to as taphonomy. This branch of science is important in helping us to understand the environmental and cultural contexts of human history. It does this by examining how the remains of organisms from the past were preserved, transported and changed. In this context, the role of gastropods is twofold: firstly, they play an important role in the reconstruction of the natural environment, and secondly, they are important in the identification of traces of human activity. Nevertheless, the potential information these remains hold has been overlooked for a considerable period, with interpretations frequently based on superficial observations or incomplete methodologies. A significant challenge encountered during the archaeological evaluation of gastropods pertains to the ambiguity surrounding the provenance of their remains, namely whether they were transported to the site by cultural or natural processes. This distinction forms the basis of taphonomic analyses (Bobrowsky, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1984\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn his comprehensive review (1984), Bobrowsky provides a detailed account of the place of gastropods in archaeomalacology, their historical development and methodological problems. In particular, four main ways in which they appear in the archaeological record are highlighted: as a food source, through natural populations, by accidental transport, or for cultural purposes (e.g. as ornaments). These findings imply that taphonomic analyses should be considered not only in a physical but also in a cultural context. These findings imply that taphonomic analyses should be considered not only in a physical but also in a cultural context. Furthermore, the preservation potential of terrestrial and freshwater gastropods may vary due to their exposure to different postmortem processes, necessitating the refinement of taphonomic analyses on a species-by-species basis.\u003c/p\u003e\u003cp\u003eAlthough the influence of invertebrates has long played a secondary role in archaeological taphonomy, it has recently been established that molluscs such as terrestrial gastropods can interact with skeletal material and leave morphologically distinguishable traces on the bone surface (Trueman \u0026amp; Martill, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). These include superficial pitting, irregular abrasions, linear gnaw marks and microscopic dissolution patterns. However, studies that systematically identify and taxonomically classify these types of scars are very limited. This has led to the attribution of these scars to other factors (microbial decomposition, root scars, erosion, etc.).\u003c/p\u003e\u003cp\u003eIn the field of gastropod taphonomy, Galvao et al. (2015) conducted a study on the evaluation of gastropod traces in forensic medicine. The study commenced with the discovery of a human skeleton buried for a period of five years in Bras\u0026iacute;lia, the capital city of Brazil. A total of 20 snail shells belonging to the Allopeas micra species were found inside the femur and humerus bones. The observation that all of the shells were found to be in the same developmental stage indicates the likelihood that they belonged to the same generation and entered the bone at approximately the same time. This finding suggests that the snails may have been actively digging into the soil and entering the bones, where they would have continued to live but eventually died. The study emphasises that the snails do not directly intervene in the body tissue, but adapt to the ecological changes in the burial environment.\u003c/p\u003e\u003cp\u003eThe objective of this study is to analyse the molluscan taphonomic traces present on the skeletal material of an individual unearthed during excavations at the ancient city of Nysa in Aydin province, western Turkey. The graves are located within a necropolis that dates to the 11th-12th century, and the soil structure is characterised by its dry, fine-grained texture and relative paucity of organic matter. The gastropods \u003cem\u003eCornu aspersum\u003c/em\u003e (=\u0026thinsp;\u003cem\u003eHelix aspersa\u003c/em\u003e), \u003cem\u003eOxychilus camelinus\u003c/em\u003e and \u003cem\u003eCaracollina lenticula\u003c/em\u003e, which were found at the same level as the skeleton of the studied individual, represent both the grave microfauna and are considered as possible perpetrators of the gnawing and superficial burrowing marks observed on the skeleton. In consideration of the ecological behaviour exhibited by these species, the inactivation of \u003cem\u003eCornu aspersum\u003c/em\u003e during both summer and winter months offers an indirect indication that the sepulchre was most likely accessed during the spring or autumn seasons, periods that are characterised by elevated levels of humidity and mild temperatures (Dahirel et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Yom-Tov, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1971\u003c/span\u003e). Consequently, this study not only documents a rare taphonomic phenomenon, but also has the potential to comment on the timing of burials and the biota of the burial environment.\u003c/p\u003e"},{"header":"2. Materials and Methods","content":"\u003cp\u003eThe skeletal material examined in this study was recovered from a grave dating to the 11th-12th century during archaeological excavations at the ancient city of Nysa in the province of Aydin in western Turkey. The grave is located in the eastern part of the necropolis area, in a stratigraphic level with a dry and fine-grained soil structure, and the burials are individual and primary. Despite the partial absence of the skeletons of the individuals exhumed from the grave, the majority of the cranial and post-cranial elements are preserved.\u003c/p\u003e\u003cp\u003eThe taphonomic marks observed on the skeletal material were documented primarily by macroscopic methods, with the naked eye and using a hand magnifier (10\u0026times;). The presence of linear and irregular grooves, pitting, and superficial abrasions on the bone surface was meticulously catalogued and categorised in accordance with the taphonomic identification system proposed by Fern\u0026aacute;ndez-Jalvo and Andrews (\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). The anatomical regions in which the lesions were found were meticulously documented, and their morphological characteristics were measured and photographed. For micro-morphological evaluation, a number of lesions were analysed under a stereo microscope (40x), with the bone surface structure being observed in detail.\u003c/p\u003e\u003cp\u003eIn addition, the skeletal remains of three distinct terrestrial gastropod species were documented within the graves at the level at which the skeleton was discovered. The species under consideration are \u003cem\u003eCornu aspersum\u003c/em\u003e (=\u0026thinsp;\u003cem\u003eHelix aspersa\u003c/em\u003e), \u003cem\u003eOxychilus camelinus\u003c/em\u003e and \u003cem\u003eCaracollina lenticula\u003c/em\u003e. In the identification of species, local malacological sources and ecological species identifiers were utilised (Yildirim et al., 2006; Schileyko, \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). The biological potential, morphological structure and feeding behaviour of these species, which may form skeletal marks, were evaluated in the light of the literature. The findings were evaluated in comparison with the relevant taphonomic literature and interpreted by distinguishing them from other possible taphonomic factors (root traces, microorganism decomposition, mechanical abrasion).\u003c/p\u003e"},{"header":"3. Results","content":"\u003cp\u003eThe taphonomic scars identified in the study are considered to be of molluscan origin. These were found on various bones belonging to different individuals. The morphology, distribution and size of the marks exhibited significant inter-individual variability. In all specimens, the marks were found to be superficial gnawing, pitting, burrowing and erosion-like. The potential effects of the identified species Cornu aspersum, \u003cem\u003eOxychilus camelinus\u003c/em\u003e and \u003cem\u003eCaracollina lenticula\u003c/em\u003e were taken into account in the evaluation of these scars.\u003c/p\u003e\u003cp\u003eOn the cranium of one individual, taphonomic scars consisting of small pits and punctures with rounded and irregular edges, measuring 1\u0026ndash;2 mm in diameter, were found on both the ectocranial and endocranial surfaces. A more comprehensive documentation of these scars was conducted on the inner surfaces of the frontal and parietal bones. The marks are deep enough to penetrate the cortical bone rather than being superficial. Irregular erosion-like abrasions were also observed on the ectocranial surface in the vicinity of some of the scars. These findings indicate long-term biological activity of molluscs in a humid environment.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eLarge and distinct scars were discovered on the right tibia of a different individual, especially in the periosteal region. Their width is approximately 2\u0026ndash;3 millimetres, and they are arranged in linear rows along the surface. Each groove manifests a round or oval morphology, meticulously organised such that it is subtly depressed into the bone surface. The sequential arrangement of these tracks is consistent with the regular gnawing of bone by a gastropod while it moves in a specific orientation. Microscopic analyses of the tracks also documented surface softening and absorption.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eTaphonomic scars of a similar nature, albeit more superficial and irregularly distributed, were observed on the ribs of another individual. The scars are particularly concentrated on the posterior rib segments and manifest as shallow, pitted grooves with a diameter of 1\u0026ndash;3 mm. In some cases, the scars were found to be in close proximity to each other, and upon closer inspection, it was observed that these scars had merged to form larger superficial abrasions. These morphological structures suggest that gastropods were concentrated in a specific area and probably had prolonged contact.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eOn the scapula bone of an individual subject, several pits with irregular edges and measuring 2\u0026ndash;3 mm in diameter were found on the surface. In the grave from which this bone was recovered, a gastropod shell was documented in situ next to the scapula. The shell is morphologically well preserved, and a taxonomic examination revealed that the grave of this individual belongs to the species Oxychilus camelinus. This provides a robust environmental context in which the traces can be directly associated with gastropod biological activity.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eGeneral Characteristics of the Findings of Tafonomy Formations\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"5\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBones\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eThrace Type\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSize (mm)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eSurface\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eInterpretation\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSkull\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003ePerforation (pits)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026ndash;2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eEndo/Ecto\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eProgressive biological wear\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTibia\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eSequential Cavity\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u0026ndash;3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePeriosteal\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eRegular Gastropod Gnawing\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eThe Rib\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eSuperficial Pit\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026ndash;3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePlevnal\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eIrregular Activity\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eScapula\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAdherent Shell\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u0026ndash;3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eLateral Surface\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eSpatial relationship with shell\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e"},{"header":"4. Discussion","content":"\u003cp\u003eThe spatial correlation of the taphonomic traces observed on the human skeletons obtained from the graves in this study with the terrestrial gastropod species found at the grave levels suggests that these traces may be of biological origin. The species under scrutiny, \u003cem\u003eCornu aspersum\u003c/em\u003e (=\u0026thinsp;\u003cem\u003eHelix aspersa\u003c/em\u003e), \u003cem\u003eOxychilus camelinus\u003c/em\u003e and \u003cem\u003eCaracollina lenticula\u003c/em\u003e, offer highly significant data within this context with regard to their ecological characteristics.\u003c/p\u003e\u003cp\u003e\u003cem\u003eCornu aspersum\u003c/em\u003e is a species that is particularly prevalent in humid and temperate regions, and it is characteristic of the Mediterranean climate. It has been documented to occur naturally on the western and southern coasts of Turkey (Herbert, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Yıldırım et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). In addition to feeding on plant material, it has been observed that the species tends to gnaw bony surfaces to meet its calcium needs (Trueman \u0026amp; Martill, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). It is evident that this species exhibits heightened activity during the spring and autumn seasons, entering periods of estivation and hibernation during summer drought and winter cold (Charrier \u0026amp; Daguzan, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1980\u003c/span\u003e). This behavioural pattern lends support to the hypothesis that the gnaw marks observed on the tibia and cranium occurred during the active seasons and that individuals were most likely buried in spring or autumn. The regularly arranged burrow marks, especially on the tibia, are consistent with the oriented gnawing behaviour exhibited by this species.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003eOxychilus camelinus\u003c/em\u003e is another species of land snail found in western and southeastern Turkey, and is adapted to moist and shaded areas (Heller et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). The feeding of this species on microbial film and decaying organic matter has been demonstrated to limit its capacity to directly damage the bone surface. However, it may support the bioerosion process occurring around the bone. The irregular and shallow pits, particularly evident on the rib bones, may be attributable to the indirect contribution of this species, a phenomenon that can be attributed to the microhabitat effect.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003eCaracollina lenticula\u003c/em\u003e, a smaller species, inhabits rocky and moist areas, especially in the Aegean Region. It has been observed that the organism feeds on organic sediments and layers of microorganisms. This observation indicates the presence of intense humidity in the burial environment. The documentation of this specimen alongside \u003cem\u003eCornu aspersum\u003c/em\u003e bark, particularly in the context of the grave where the scapula bone was unearthed, provides a direct spatial context for the origin of the traces observed on the bone. In consideration of the life cycles of the aforementioned species, which average at between two and five years, the traces observed on the bone surface appear to be the result of protracted biological activity within the inhumation environment.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eFurthermore, the seasonal behaviour exhibited by the species offers indirect insights into the timing of burial. The observation that \u003cem\u003eCornu aspersum\u003c/em\u003e demonstrates the highest levels of activity during both spring and autumn supports the hypothesis that burials may have occurred during these seasons. The observations made on the endocranial surface indicate that molluscs gained entry to the cranial cavities and established long-term biological contact.\u003c/p\u003e\u003cp\u003eThe active involvement of these three species in the microecology of the grave provides information on taphonomic aspects, as well as the moisture balance, soil structure and seasonal climatic conditions of the grave environment. The rarity of the documentation of gastropod-derived taphonomic traces on human skeletons in the literature (see Fern\u0026aacute;ndez-Jalvo \u0026amp; Andrews, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) renders this study a significant exemplar within this field. In consideration of the morphology, dietary biology and environmental sensitivity of gastropod species, it is evident that these species may have played an active role in intra-grave biological processes.\u003c/p\u003e\u003cp\u003eThe taphonomic traces documented in this study are most likely the result of biological activity by molluscs, given their morphological structure, distribution on bone, and spatial relationships with gastropod species found in the grave contents. In particular, the gnawing behaviour of species such as \u003cem\u003eCornu aspersum\u003c/em\u003e on hard surfaces to meet their calcium carbonate needs has been previously observed in both natural and experimental environments (Trueman \u0026amp; Martill, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Yom-Tov, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1971\u003c/span\u003e). The probability of biological interactions in graves increases during the spring and autumn months when this species is active outside the estivation and hibernation periods.\u003c/p\u003e\u003cp\u003eThe presence of pore and pit marks on both the endocranial and ectocranial surfaces of the skull is a distinctive feature that has not been previously documented in the literature. In particular, in the endocranial regions, these marks indicate that the organisms had a long-term presence in the skeletal cavities and had access to both surfaces of the bone. Furthermore, the sequential burrow marks on the tibia indicate that \u003cem\u003eCornu aspersum\u003c/em\u003e gnawed while moving in a controlled orientation. This regularity lends support to the hypothesis that the tracks are not random, but rather based on biological behaviour.\u003c/p\u003e\u003cp\u003eIt is important to note the potential for such traces to be misinterpreted as other taphonomic factors, including but not limited to root action, insect activity, or chemical dissolution. However, the presence of a gastropod shell alongside a scapula bone in an in situ grave suggests that this effect may not be accidental, but rather the result of a systematic interaction specific to the grave microenvironment. In a similar vein, Marchiafava et al. (2015) conducted an experimental study that demonstrated the presence of molluscs in graves can leave marks on bone surfaces. However, the fact that the traces detected in this study exhibit a more intense and different distribution in terms of diameter and depth reveals the uniqueness of the Nysa sample.\u003c/p\u003e\u003cp\u003eIn the existing literature, the taphonomic effects of molluscs are predominantly discussed in the context of animal bones or prehistoric contexts, while their effects on human skeletons are seldom documented (Fern\u0026aacute;ndez-Jalvo \u0026amp; Andrews, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Consequently, the data presented here are significant for two reasons. Firstly, they serve to document a new group of agents in the field of archaeological taphonomy. Secondly, they facilitate a more profound comprehension of inhumation biological processes.\u003c/p\u003e"},{"header":"5. Conclusions","content":"\u003cp\u003eThis study revealed that the taphonomic marks observed on human skeletons recovered from 11th-12th century tombs of the ancient city of Nysa may be related to the biological activities of terrestrial gastropods. The species Cornu aspersum, \u003cem\u003eOxychilus camelinus\u003c/em\u003e and \u003cem\u003eCaracollina lenticula\u003c/em\u003e were identified as potential factors in the formation of these marks, and were evaluated in terms of both their morphological and ecological characteristics. In particular, the coincidence between the feeding behaviour and seasonal activities of the species with the punctures, sequential burrows and superficial erosion-like scars seen on the bone surface indicates that these scars may be of gastropod origin.\u003c/p\u003e\u003cp\u003eThe presence of traces on various bone regions, including the skull, tibia, ribs, and scapula, has enabled the formulation of inferences concerning not only the biological processes within the grave, but also the period of burial, the environmental moisture balance, and the post-burial microfaunal dynamics. \u003cem\u003eCornu aspersum\u003c/em\u003e has been observed to exhibit heightened activity during the spring and autumn months, thereby supporting the hypothesis that individuals may have been interred during these periods.\u003c/p\u003e\u003cp\u003eThe fact that the traces left by gastropods on skeletal material in archaeological contexts are often overlooked increases the importance of this study. In this context, the study introduces a novel perspective to research on both human taphonomy and funerary microfauna, thereby drawing attention to the taphonomic role of terrestrial molluscs. In the future, further studies supported by experimental taphonomy, microscopic analyses and environmental data will serve to reinforce the accuracy of these findings and provide more refined results in taphonomic interpretations.\u003c/p\u003e"},{"header":"6. Limitations","content":"\u003cp\u003eDespite the fact that the present study sets out to establish the relationship between molluscan taphonomic tracks and their possible perpetrators, it is important to consider the limitations of the study. Firstly, the fact that the traces were evaluated solely on the basis of morphological observation and contextual association limits the precise establishment of a cause-effect relationship. In particular, microscopic or chemical trace analyses were not performed in order to determine the gastropod origin of the pits and abrasions seen on the bone surfaces, and compositional characterisation of the traces was not possible.\u003c/p\u003e\u003cp\u003eFurthermore, given the potential for other biological agents (e.g. microorganisms, insects, root systems) present in the grave contents to leave similar traces on the bones, the necessity for additional comparative samples is apparent, with the aim of excluding alternative taphonomic factors. In this context, the hypothesis that the present traces have a molluscan origin is regarded as the most probable scenario in view of the extant data; nevertheless, it is acknowledged that alternative explanations cannot be wholly excluded.\u003c/p\u003e\u003cp\u003eIn subsequent studies, experimental taphonomy research in controlled laboratory settings can be utilised to more systematically document how different gastropod species leave marks on the bone surface. Furthermore, the employment of advanced imaging techniques, such as scanning electron microscopy (SEM) or micro-CT scans, facilitates a more detailed analysis of the micromorphology of the tracks. Furthermore, the impact of environmental factors associated with the burial environment, including soil chemistry, moisture content and seasonal variability, on trace formation will be a pivotal area of focus in future research endeavours.\u003c/p\u003e\u003cp\u003eFinally, the under-documentation of such traces in the literature necessitates more careful monitoring of the taphonomic impact of molluscs in archaeological excavations and systematic recording of microfauna data.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eAll work belongs to the responsible author.\u003c/p\u003e\u003ch2\u003eAcknowledgements\u003c/h2\u003e\u003cp\u003eI would like to express my gratitude to Professor \u0026Uuml;mit Kebap\u0026ccedil;ı for his assistance with the identification of gastropod species and Professor Serdar Hakan \u0026Ouml;ztaner, head of the Nysa Excavation, for his support.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eBehrensmeyer, A. K. (1978). Taphonomic and Ecologic Information from Bone Weathering. \u003cem\u003ePaleobiology\u003c/em\u003e, 4(2), 150-162. Doi:10.1017/S0094837300005820\u003c/li\u003e\n\u003cli\u003eBobrowsky, P. T. (1984). The History and Science of Gastropods in Archaeology. \u003cem\u003eAmerican Antiquity\u003c/em\u003e, 49(1), 77\u0026mdash;93. https://doi.org/10.2307/280513\u003c/li\u003e\n\u003cli\u003eCharrier, M., \u0026amp; Daguzan, J. (1980). Consommation alimentaire, production et bilan \u0026eacute;nerg\u0026eacute;tique chez \u003cem\u003eHelix aspersa\u003c/em\u003e m\u0026uuml;ller (gast\u0026eacute;ropode pulmon\u0026eacute; terrestre) [Food consumption, production and energy evaluation in \u003cem\u003eHelix aspersa\u003c/em\u003e m\u0026uuml;ller (a terrestrial pulmonated gasteropod)]. \u003cem\u003eAnnales de la nutrition et de l\u0026apos;alimentation\u003c/em\u003e, \u003cem\u003e34\u003c/em\u003e(1), 147\u0026ndash;166.\u003c/li\u003e\n\u003cli\u003eDahirel, M., Olivier, E., Guiller, A., Martin, M.-C., Madec, L. and Ansart, A. (2015), Movement propensity and ability correlate with ecological specialization in European land snails: comparative analysis of a dispersal syndrome. \u003cem\u003eJ Anim Ecol\u003c/em\u003e, 84: 228-238. https://doi.org/10.1111/1365-2656.12276\u003c/li\u003e\n\u003cli\u003eFern\u0026aacute;ndez-Jalvo, Y., \u0026amp; Andrews, P. (2016). Atlas of Taphonomic Identifications. 1001+ Images of Fossil and Recent Mammal Bone Modification. The Netherlands: Springer. https://doi.org/10.1007/978-94-017-7432-1\u003c/li\u003e\n\u003cli\u003eGalv\u0026atilde;o, M. F., Pujol-Luz, J. R., de Assis Pujol-Luz, C. V., de Rosa, C. T., Simone, L. R., B\u0026aacute;o, S. N., Barros-Cordeiro, K. B., Pessoa, L., \u0026amp; Bissacot, G. (2015). Shells and Bones: A Forensic Medicine Study of the Association of Terrestrial Snail Allopeas micra with Buried Human Remains in Brazil. \u003cem\u003eJournal of forensic sciences\u003c/em\u003e, \u003cem\u003e60\u003c/em\u003e(5), 1369\u0026ndash;1372. https://doi.org/10.1111/1556-4029.12882\u003c/li\u003e\n\u003cli\u003eHeller, J., Sivan, N. and Hodgson, A.N. (1997), Reproductive biology and population dynamics of an ovoviviparous land snail, \u003cem\u003eLauria cylindracea\u003c/em\u003e (Pupillidae). \u003cem\u003eJournal of Zoology\u003c/em\u003e, 243: 263-280. https://doi.org/10.1111/j.1469-7998.1997.tb02781.x\u003c/li\u003e\n\u003cli\u003eHerbert, D.G. (2010). The introduced terrestrial mollusca of South Africa. SANBI, Biodiversity Series, 15\u003c/li\u003e\n\u003cli\u003eLyman, R. L. (1994). \u003cem\u003eVertebrate Taphonomy. \u003c/em\u003eCambridge: Cambridge University Press.\u003c/li\u003e\n\u003cli\u003eSchileyko, A. A. (2003). Treatise on Recent terrestrial pulmonate molluscs. Part 10. Ariophantidae, Ostracolethidae, Ryssotidae, Milacidae, Dyakiidae, Staffordiidae, Gastrodontidae, Zonitidae, Daudebardiidae, Parmacellidae. \u003cem\u003eRuthenica.\u003c/em\u003e Supplement 2: 1309-1466.\u003c/li\u003e\n\u003cli\u003eTrueman, C.N. and Martill, D.M. (2002), The long\u0026ndash;term survival of bone: the role of bioerosion. Archaeometry, 44: 371-382. https://doi.org/10.1111/1475-4754.t01-1-00070\u003c/li\u003e\n\u003cli\u003eYom-Tov, T. (1971). Body Temperature And Lıght Reflectance In Two Desert Snaıls, \u003cem\u003eJournal of Molluscan Studies\u003c/em\u003e, Volume 39, Issue 4, 319\u0026ndash;326, https://doi.org/10.1093/oxfordjournals.mollus.a065111\u003c/li\u003e\n\u003cli\u003eYıldırım, M.Z., Koca S.B., Kebap\u0026ccedil;ı, \u0026Uuml;. (2006). Supplement to the Prosobranchia (Mollusca: Gastropoda) Fauna of Fresh and Brackish Waters of Turkey. \u003cem\u003eTurkish Journal of Zoology\u003c/em\u003e. 30: 197-204. \u003cem\u003e(PDF) The gastropod fauna and their abundance, and some physicochemical parameters of Lake G\u0026ouml;lbaşı (Hatay, Turkey)\u003c/em\u003e.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Taphonomy, Bioarchaeology, Nysa, Burial, Forensic Archaeology","lastPublishedDoi":"10.21203/rs.3.rs-7032422/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7032422/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThis study investigates mollusc-induced taphonomic modifications on human skeletal remains unearthed from 11th\u0026ndash;12th century burials at the ancient city of Nysa (Aydın, western Turkey). Taphonomic alterations\u0026mdash;such as pitting, linear grooves, and superficial abrasions\u0026mdash;were identified on cranial, tibial, rib, and scapular bones. These modifications were evaluated in connection with three terrestrial gastropod species found in situ: \u003cem\u003eCornu aspersum\u003c/em\u003e (=\u0026thinsp;\u003cem\u003eHelix aspersa\u003c/em\u003e), \u003cem\u003eOxychilus camelinus\u003c/em\u003e, and \u003cem\u003eCaracollina lenticula\u003c/em\u003e. Macroscopic and stereomicroscopic observations revealed distinct morphological patterns consistent with biological activity, particularly the gnawing and burrowing behavior typical of \u003cem\u003eCornu aspersum\u003c/em\u003e. Endocranial and ectocranial surfaces of cranial bones exhibited small perforations (1\u0026ndash;2 mm), while the tibiae showed linear, sequential cavities (2\u0026ndash;3 mm), indicating prolonged and directed contact. The spatial association of gastropod shells and modified bones, especially the scapula, strengthens the hypothesis of direct biological interaction. Furthermore, ecological traits and seasonal activity cycles of the gastropods support the inference that the burials may have occurred during spring or autumn, when gastropod activity peaks. This study documents a rarely addressed taphonomic phenomenon and emphasizes the interpretive potential of gastropod-bone interactions for reconstructing burial conditions, microenvironmental dynamics, and seasonality. By integrating malacological, archaeological, and taphonomic evidence, the research contributes to a more nuanced understanding of inhumation processes and highlights the need to consider invertebrate activity as a significant factor in post-mortem bone alteration.\u003c/p\u003e","manuscriptTitle":"Terrestrial Gastropod Traces on Medieval Human Remains: Taphonomic Evidence from Nysa in Western Anatolia","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-07-25 04:59:10","doi":"10.21203/rs.3.rs-7032422/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"2fc788dc-d91c-4afd-98c5-65b84d351991","owner":[],"postedDate":"July 25th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2025-11-20T20:38:08+00:00","versionOfRecord":[],"versionCreatedAt":"2025-07-25 04:59:10","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7032422","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7032422","identity":"rs-7032422","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}