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Article

From Prey to Pattern: Integrating Faunal and Behavioural Evidence of Neanderthal Subsistence at Fumane Cave (Unit A9), Northern Italy

by
Kalangi Rodrigo
1,2,*,
Nicola Nannini
3,
Vittorio Facincani
1,
Matteo De Lorenzi
1 and
Marco Peresani
1,4,*
1
Dipartimento di Studi Umanistici, Sezione di Scienze Preistoriche e Antropologiche, Università di Ferrara, Corso Ercole I d’Este 32, 44121 Ferrara, Italy
2
Doctoral Training Centre, University of Oxford, 1-4 Keble Rd, Oxford OX1 3NP, UK
3
MUSE—Museo Delle Scienze, Corso del Lavoro e della Scienza 3, 38123 Trento, Italy
4
Istituto di Geologia Ambientale e Geoingegneria, Consiglio Nazionale delle Ricerche, Piazza della Scienza 1, 20126 Milano, Italy
*
Authors to whom correspondence should be addressed.
Quaternary 2026, 9(1), 14; https://doi.org/10.3390/quat9010014
Submission received: 25 November 2025 / Revised: 25 January 2026 / Accepted: 5 February 2026 / Published: 9 February 2026
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Abstract

This study presents a zooarchaeological and taphonomic analysis of the previously unstudied component of the Mousterian faunal assemblage from Unit A9 at Grotta di Fumane (northeastern Italy), offering refined insights into Neanderthal subsistence behaviour during Marine Isotope Stage 3. Building on the previously published analysis of the principal portion of the assemblage, the new data reaffirm a subsistence strategy focused on selective transport and intensive on-site processing of high-utility carcass components. The ungulate assemblage—dominated by Cervus elaphus and Capreolus capreolus, with additional contributions from Rupicapra rupicapra and Capra ibex—is characterised by the dominance of hindlimb elements, moderate cranial representation, and a pronounced scarcity of axial remains. These patterns indicate that carcass reduction commenced at kill sites, where low-yield trunk segments were removed, while high-nutritional-value limb portions were preferentially transported to the cave for secondary processing. Taphonomic indicators, including abundant cut marks, percussion notches, and extensive bone fragmentation, demonstrate systematic defleshing, marrow extraction, and possible grease rendering within the cave, activities that were spatially associated with combustion features. Occasional cranial transport suggests targeted acquisition of high-fat tissues such as brains and tongue, behaviour consistent with cold-climate optimisation strategies documented in both ethnographic and experimental contexts. Collectively, the evidence indicates that Unit A9 served as a residential locus embedded within a logistically organised mobility system, where carcass processing, resource exploitation, and lithic activities were closely integrated. These findings reinforce the broader picture of late Neanderthals as adaptable and behaviourally sophisticated foragers capable of strategic planning and efficient exploitation of ungulate prey within the dynamic environments of northern Italy.

1. Introduction

This study presents a zooarchaeological and taphonomic analysis of a new bone assemblage from Grotta di Fumane, the Mousterian Unit A9, where discoid manufacturing lithic industries are extensively reported [1,2,3]. The aim of this research is to provide a complete picture of the faunal assemblage by thoroughly examining the substantial bone assemblage recovered from the innermost excavated area of the site, as this extensive collection has the potential to offer valuable insights into the subsistence strategies and foraging behaviours of the Neanderthal inhabitants.
The Grotta di Fumane, situated in the foothills of the Venetian Prealps in the western Lessini Mountains, north-eastern Italy (Figure 1A), has been extensively investigated over two decades. This region characterised by a complex network of active fault systems that have shaped the local topography over time [4], which provides a unique opportunity to explore Neanderthal foraging behaviour during the late Middle Palaeolithic period [5]. This period is characterised by extensive cultural replacements, as indicated by diverse archaeological signatures, such as technological innovations that are the product of adaptive strategies and changes in dietary habits [6,7]. The Mousterian Unit A9 preserves a well-defined cultural layer [8,9] that can provide additional insights and contribute to the understanding of these topics through the study of its faunal assemblage.
Human occupation at Fumane Unit A9 is attested to by numerous faunal remains, lithic artefacts, and combustion features [5] contained in a 15–20 cm thick stratigraphic complex characterised by overlapping thin layers and lenses. The composition includes frost-shattered stones, aeolian silt and sands, and dark sediments, all extensively distributed due to intense human activities. Topographical mapping has identified intact or fragmented bones, teeth, and identifiable fragments ≥ 5 cm, while smaller fragments, micromammal and small avifaunal bones were recovered through wet sieving [1].

2. Materials and Methods

Zooarchaeological and Taphonomical Frame of Reference

The faunal material discussed here represents the portion of the A9 assemblage that was unexamined until this study, while the rest had already been analysed and published in 2014 [1]. Zooarchaeological analysis was conducted on the entire faunal assemblage recovered from 1 m2 squares 137, 147, 157, 158, 167, and 168, totalling 6 m2 excavated in the innermost explored area beyond the cave entrance (Figure 1B). To determine anatomical and taxonomical classifications, first, the assemblage was classified following the same dimensional division method used in the previous study (0–1 cm, 1–2 cm, 2–3 cm, 3–4 cm, 4–5 cm, 5 cm). Additionally, burned and calcined bones were segregated from unburned specimens. Taxonomic and anatomical identifications in this study were based on established zooarchaeological comparative alpine and archaeological collections of the University of Ferrara and the Laboratory of Prehistory and Zooarchaeology of MUSE, the Museum of Science, and a series of reputable publications [10,11,12,13,14]. Microscopic analyses of bone surfaces were aided by a Leica S6D Greenough stereomicroscope with 0.75–70x magnification range. The studied material was recorded in a database specifically catered for the study’s aims.
Both identified remains, and those deemed unidentifiable based on morphological or size characteristics were categorised into three mammal body-size classes, following the methodology of Bunn et al. [15]: I—small (i.e., Lepus sp. and other lagomorphs, Mustelidae, Vulpes vulpes); II—medium (i.e., Capreolus capreolus, Rupicapra rupicapra, Capra ibex, Canis lupus); and III—large (i.e., Cervus elaphus, Ursidae, Megaloceros giganteus, Bovinae). When species determination is not possible, specimens were identified by their family (e.g., Cervidae, Caprinae) or genus (e.g., Capra/Cervus, Rupicapra/Capreolus, Bos/Bison sp.) level.
According to modern animal birth criteria, the age at death and the state of epiphysis fusion were determined using tooth eruption, replacement sequences, and dental wear. Priority was given to ungulates, such as Capreolus capreolus [16], Cervus elaphus [17,18,19], and bovids [20,21,22], due to preservation bias.
All taxonomically identified specimens, including long bone fragments, were considered in the analysis of skeletal part profiles. NISPs (Number of Identified Specimens) [23] and MNIs (Minimum Number of Individuals) [24] indices were used in order to evaluate species abundance. MNIs were estimated considering all skeletal elements, including both bones and teeth, with a particular emphasis on the tibia for ungulates and teeth for carnivores (if any). MNEs (Minimum Number of Elements) were calculated by selecting the anatomical zone with the highest representation of >50%, which was further combined with side and fusion data for each specific element.
The identification of taphonomic alterations, whether organic (microorganism, animal, plant, etc.) or inorganic and mechanical (manganese stains, weathering, water circulation or sediment mobilisation, sediment compaction, etc.), was based on previously published criteria [12,25,26,27,28,29,30,31,32,33,34,35]. The identification of anthropic modification was based on examining the position, type, and orientation of linear marks. Incisions and scraping marks were identified as types of cutmarks [28,30,36,37,38]. An analysis of cutmarks was conducted in order to infer possible actions carried out on the bone, such as skinning, disarticulation, defleshing, tendons removal and periosteum removal [39]. The analysis considered the number of striations, their location and distribution, and orientation. Furthermore, the presence of chop marks was recorded. Chop marks refer to broad and relatively short linear depressions with a V-shaped cross section [12,29]. Anthropic traces resulting from bone breakage for extracting marrow were identified and categorised into two types: percussion marks and impact flakes. Percussion marks take the form of semi-circular cavities located on the fracture edges, with corresponding negative flake scars. Impact flakes are positive flakes of the percussion notches that result from breaking the diaphysis [32,33,40,41,42]. Deliberate bone fracturing to reach marrow was identified based on diagnostic green-bone fracture features at shaft extremities, including spiral/oblique breaks, smooth surfaces, and acute fracture angles indicative of percussive breakage [29,34,43,44,45,46]. Every anatomical element (over 3 cm long) was examined both macroscopically and microscopically to identify surface alterations. The degree of combustion was evaluated by the methodology developed by Stiner et al. [47], whereby bones are assigned to combustion stages (unburned, carbonised, partially calcined, fully calcined) based on macroscopic colour and surface alterations. Assessing the skeletal representation of various species and the survival rates of bones was based on fragmentation indices methodologies developed by Binford [30], Brain [31], and Lyman [12].
Habitat preferences of species in the ungulate assemblage suggest two habitat types [48,49]. Ungulates such as moose (Alces alces), giant deer (Megaloceros giganteus), red deer (Cervus elaphus), roedeer (Capreolus capreolus), and wild boar (Sus scrofa) represent a mixed habitat, such as grasslands, wetlands and forests of the lower elevations, while mountainous habitats were preferred by ibex (Capra ibex) and chamois (Rupicapra rupicapra), in higher elevations than the former.

3. Results

3.1. Zooarchaeology and Taphonomy

Out of a total of 22, 990 bone fragments (NR) chosen for analysis, 1738 (7.5%) (NISPs) were sufficiently preserved to allow for morphological assessment. Of these, 424 specimens (24.4%) were successfully identified to species or genus level (Table 1). Natural modifications in the assemblage primarily consist of manganese stains (35.6%), followed by root marks (25.8%). Weathering-related micro fissures (4.3%) and trampling (5.1%) are comparatively low, while other modifications, including exfoliations (1.4%), carbonate concentrations (0.7%), carnivore marks (1.7%), rodent gnawing marks (1.1%), and corrosion (0.3%), each account for less than 6% of the total. Burned fragments account for 15% of the total, while calcined fragments make up approximately 9% of the total assemblage (Figure 2). Carnivore and rodent contributions to the accumulation are minimal, collectively do not exceed 3% of the total assemblage, suggesting that carnivores may had secondary access to the assemblage, but did not make a huge impact on the bone accumulation.
Ungulates represent nearly 96% of the total assemblage, primarily through limb bones (Figure 3). Red deer and roe deer are the most common species, indicating a sparse forest environment [50,51,52], though the relatively low presence of giant deer and moose is indicative of a cold-temperate environment with occasional access to open grazing areas. These ecological conditions are consistent with those found in the vicinity of the site, which could also serve as a refuge for species such as chamois and ibex. The high frequency of cervids aligns with faunal records from Mousterian contexts globally [53,54,55,56,57]. In the preceding layers (A11-10) at Fumane, there is a noticeable increase in these species, with a corresponding decline in species adapted to open environments [1].
Hunting and foraging practices appear to be biased towards forested environments, as reflected in the taxonomic composition of the assemblage. This suggests a focus on lowland areas and valley bottoms. Larger prey such as moose and giant deer likely had lower encounter frequencies and hunting success rates than red deer and roe deer, owing to their lower population densities, more restricted habitat use, and higher pursuit and handling costs [58,59,60,61]. However, these larger species may have been more successfully hunted than smaller prey in higher altitudes.
A range of species can be identified among the carnivore remains. The presence of the red fox (Vulpes vulpes) and ursids (genus Ursus) indicates a preference for forested habitats [62]. Cut marks on a complete first phalanx of a brown bear (Ursus arctos) are the only human-induced modification found in the carnivore assemblage. The cut marks on this bone suggest skinning, possibly related to the removal of fur for utilitarian or subsistence purposes.

3.2. Exploitation of Carcasses

Anthropic marks, indicative of butchering activities, are observed on the surfaces and fresh fracture edges of bones from all species of ungulates (Table 2). The presence of cranial elements suggests that large ungulates, such as red deer, were processed differently from smaller prey species, including roe deer, chamois, and ibex. The presence of cranial elements suggests that whole carcasses were transported to the camp, rather than just the more economically advantageous meaty portions being brought to the site (Figure 4).
Cut marks made by lithic tools are abundant, with 111 marks found on ungulates (Table 2) and 86 on undetermined bone fragments. The majority of taxonomically identifiable butchered specimens are red deer, with 33 specimens bearing cut marks and percussion marks (Table 3, Figure 5 and Figure 6). There were fewer butchering marks on other commonly represented species, such as roe deer (n = 11), chamois (n = 5), and ibex (n = 3). This can be partly explained by the relatively high number of undetermined ungulate remains bearing cut marks (n = 65), suggesting that a larger proportion of these species were processed but cannot be precisely taxonomically identified.
Four bones bearing retouching impact marks (Figure 6): two are undetermined ungulates, and one each is red deer and chamois. Stigmata-like punctiform impressions, linear impressions, retouch-induced striae, and notches can be explained by the fact that the bones were used as hammers to knap flint.

3.3. Carcass Representation Index

The A9 ungulate assemblage reveals a clear and structured pattern of skeletal-part representation that provides important insights into carcass transport and butchering practices carried out by Neanderthal groups at Fumane Cave. Cranial elements account for 13.2% of the identifiable remains, most of which are isolated teeth. This moderate proportion indicates occasional rather than systematic transport of heads, likely reflecting opportunistic acquisition of the high-fat resources contained within the cranium. Axial remains comprise only 2.7% of the assemblage, a figure consistent with the near absence of vertebrae and ribs throughout the layer. The scarcity of these bulky, low-utility elements strongly suggests that primary disarticulation occurred at kill sites, where axial portions were routinely discarded prior to transport.
Appendicular elements dominate the assemblage. Forelimb bones represent 18.5% of the total, whereas hindlimb elements form the largest category at 33.5%. The marked abundance of femora, tibiae, and metatarsals reflects their high flesh and marrow yield and corresponds to a transport strategy prioritising anatomical segments with the greatest caloric return. Autopodial elements—including metapodials, phalanges, and associated carpals and tarsals—comprise 21.5% of the assemblage (Figure 4 and Figure 7). Their notable frequencies relate both to their intrinsic structural durability and to the regular transport of robust distal parts attached to major limb segments. This distribution closely parallels the pattern described for A9 by Romandini et al. [1], who likewise reported a dominance of hindlimb elements, moderate cranial representation, and minimal axial remains. Combined, the two datasets present a consistent behavioural signal: carcasses were initially reduced at kill locations, and only the nutritionally richest portions (and some lower-nutrition portions that were attached to these higher-nutrition parts) were transported to the cave. Intermediate frequencies of forelimb and cranial elements point to a flexible, yet consistently selective transport strategy designed to maximise caloric gain while minimising the labour involved in moving heavy, low-yield anatomical parts.
Evidence for intensive on-site processing—including high levels of limb fragmentation and frequent cut and percussion marks on transported elements—indicates defleshing and marrow extraction within the cave. These modifications support the interpretation of Unit A9 as a habitation area where carcass reduction was routinely completed. Overall, the anatomical and taphonomic data reflect a structured and efficient subsistence system in which Neanderthal groups employed cost-effective transport and processing behaviours.

4. Discussion

The findings of this paper are consistent with those from the previously analysed section of the Fumane A9 unit [1]. The earlier analysis covered the majority of the faunal assemblage, and the present study examines the material that had not yet been analysed. With this analysis, the study of the faunal material from Fumane cave’s A9 layer is now complete.
Post-depositional taphonomic alterations are dominated by manganese oxide staining and root etching (Figure 2), indicating prolonged exposure of the assemblage to humid sedimentary conditions. High frequencies of manganese staining are commonly associated with reducing environments and fluctuating moisture conditions within sediments [63,64,65]. The widespread occurrence of root marks further points to post-depositional interaction with vegetation and paedogenic activity, suggesting relatively stable sediments and soil formation processes following burial [66,67]. In contrast, the low incidence of weathering, exfoliation, and corrosion indicates limited subaerial exposure prior to burial and relatively rapid sediment cover, which reduced prolonged atmospheric degradation of bone surfaces [25]. Moderate frequencies of trampling abrasions are consistent with localised sediment compaction or minor post-depositional disturbance within the cave [29,68,69], while the scarcity of carnivore and rodent gnawing suggests minimal post-depositional biotic modification by scavengers [31]. Overall, the taphonomic signature is consistent with limited reworking and generally good preservation of anthropogenic surface modifications.
The zooarchaeological evidence offers a robust basis for reconstructing Neanderthal subsistence behaviour within the region. From an ecological perspective, the species present are consistent with an environment characterised by sparse forest cover interspersed with open grazing areas. Such a mosaic landscape appears to have encouraged a flexible, opportunistic approach to hunting rather than a strategy targeting specific taxa. The diversity of species represented, coupled with the presence of individuals of varying ages, supports the interpretation of a generalist subsistence strategy adapted to the environmental heterogeneity of the Lessini plateau. Comparable behavioural plasticity has been documented in units A5–A6 at Fumane and at other MIS 3 Mousterian sites in northern Italy, including Riparo Tagliente [70,71], Riparo Mezzena [72], and Grotta San Bernardino [73,74,75].
The skeletal-part representation of the A9 ungulate assemblage provides a coherent picture of carcass transport and processing practices during the late Middle Palaeolithic occupation of the site. Appendicular elements dominate the assemblage, with hindlimbs accounting for 33.5% and autopodial elements for 21.5%, while axial remains are scarce overall (2.7%), with cranial elements (13.2%) and forelimbs (18.5%) occurring in more moderate proportions. This structured distribution mirrors the pattern reported in the earlier A9 study [1] and indicates consistent, economically rational transport strategies shaped by prey size, terrain constraints, and logistical considerations such as distance from kill sites [30,76,77].
Axial elements suggest that carcass processing typically began at kill locations, where bulky, low-utility components were discarded [30,77,78,79,80,81,82]. This behaviour aligns with established models of transport optimisation [30,78,83], which predict that hunters negotiating complex topography, such as the Monti Lessini foothills, would prioritise the removal of high-yield, low-weight anatomical parts. The predominance of hindlimb elements—rich in flesh and marrow—supports this interpretation, as does the high frequency of anthropogenic modifications, including cut marks, percussion damage, and fragmentation indicative of marrow extraction. Taken together, the evidence points to the regular introduction of high-utility limb elements into the cave for intensive secondary processing.
Cranial representation (13.2%) reflects a more variable transport pattern. Occasional transport of skulls is consistent with the acquisition of nutritionally valuable resources such as brain, tongue, and facial musculature—packages particularly advantageous during colder climatic phases (biochemical basis, e.g., review of brain lipids; see [84,85]). Taphonomic evidence of cranial cut marks and percussion in Neanderthal contexts, such as Goyet [86], combined with zooarchaeological analysis documenting skull exploitation and selective transport [30,87], further supports this behaviour. However, the moderate frequency of cranial remains indicates that head transport was not systematic, reinforcing the interpretation that limb elements were preferentially transported over less predictable nutrient sources. This pattern parallels the earlier findings for A9 [1], which likewise suggested context-dependent decisions regarding head transport.
The relatively high proportion of autopodial elements (21.5%) merits particular attention. Although some of this representation can be attributed to the high density and consequent survivorship of distal limb bones, the recurrent presence of metapodials and phalanges suggests that entire limbs or large limb sections were frequently transported. Similar patterns have been documented at Middle Palaeolithic sites such as Bolomor Cave [88], where distal limb segments accompanied proximal ones due to natural anatomical attachment and their supplementary value for marrow, tendon, and hide processing. The A9 assemblage thus indicates that transported limb packages were not restricted solely to the most profitable components but often included a broader anatomical range.
Comparative data from other European Middle Palaeolithic contexts further situate the Fumane A9 findings. Selective limb transport at sites such as Abri du Maras (Rhône Valley) [89] and Combe-Grenal (France) [90,91] has been linked to mobility systems in which kill locations were spatially separated from residential hubs. The A9 anatomical profile conforms to this pattern, suggesting that Neanderthal groups hunted red deer and roe deer in nearby lowlands and transported high-utility body parts to the cave for processing. Combustion features and dense concentrations of fragmented limb bones at the cave entrance [1] reinforce the interpretation of A9 as a locus for regular food processing rather than short-term or task-specific use. Collectively, the anatomical representation, modification signatures, and inter-site comparisons indicate a subsistence system that maximised caloric return through coordinated transport and processing within a logistical mobility framework.
The presence of a single cut-marked bear bone, although limited in number, contributes to growing evidence for infrequent but meaningful Neanderthal exploitation of ursids in northern Italy. As noted by Romandini et al. [92], MIS 3 Mousterian sites such as Grotta San Bernardino, Grotta Rio Secco in the Carnic Prealps, and Grotta di Fumane preserve clear indicators of bear carcass processing. While sporadic, such evidence underscores the flexibility of Neanderthal foraging strategies and their willingness to exploit a wide array of available resources. Taphonomic observations reveal minimal evidence of carnivore or rodent disturbance and low levels of weathering, confirming a predominantly anthropogenic accumulation. Burned and calcined remains, representing approximately 4% of the assemblage, attest to proximity to hearths [5,9] and the close association between faunal processing and combustion activities. The combined presence of cut marks, percussion damage, and retoucher use supports the interpretation of A9 as a residential and processing area rather than a secondary deposition zone. Balanced representation of axial and appendicular remains for larger ungulates, together with intensive butchery evidence, further suggests that primary carcass processing, marrow extraction, and possibly hide preparation occurred directly within the cave. These patterns are consistent with other structured on-site faunal exploitation documented in Italian Mousterian contexts, such as Grotta de Nadale [93,94].
The faunal evidence highlights the behavioural sophistication of Neanderthal groups in the region. Subsistence practices at Fumane reflect logistical planning, broad-spectrum prey selection, and efficient utilisation of carcass resources, paralleling patterns observed in some Middle Palaeolithic contexts [95,96,97]. The integration of hunting, transport, butchery, and resource processing—supplemented by evidence for less frequent cranial exploitation, retoucher production, and controlled burning—demonstrates a high degree of cognitive flexibility and ecological awareness. Within the broader European context, the Fumane A9 assemblage aligns with an increasingly diversified Mousterian subsistence pattern during MIS 3, comparable to sites such as Teixoneres Cave (Spain) and Pié Lombard (France) [98]. Among northern Italian sites, Fumane stands out for its integrated faunal, lithic, and combustion evidence within a single stratigraphic unit, offering one of the clearest examples of a well-organised residential base from this period [9].

5. Conclusions

The faunal assemblage from Unit A9 at Grotta di Fumane provides a refined perspective on Neanderthal subsistence behaviour on the Lessini Plateau during MIS 3. Rather than reflecting purely opportunistic foraging, the anatomical distribution of ungulate remains indicates a structured and economically informed transport strategy. The strong predominance of hindlimb elements, together with the scarcity of postcranial axial remains (e.g., vertebrae and ribs) and only moderate representation of cranial elements, suggests that carcass processing typically began at kill locations, where bulky, low-utility components were discarded. High-yield portions—particularly femora, tibiae, and metatarsals—were selectively transported to the cave for intensive secondary processing. This pattern mirrors that identified in the previously analysed A9 assemblage [1] and aligns with broader Middle Palaeolithic evidence from sites such as Abri du Maras [42,89,99] and Combe-Grenal [90,91], where selective limb transport has been interpreted as a key feature of logistically organised mobility systems.
The frequent occurrence of cut marks, percussion impacts, and extensive fragmentation further indicates that A9 functioned as a residential locus where carcass reduction and marrow extraction were routinely carried out. The presence of cranial elements, though at moderate levels, points to occasional skull transport for the acquisition of nutrient-rich tissues such as brain and tongue—behaviours consistent with ethnographic and experimental models of fat-oriented transport during cold climatic phases [30,78]. Additional evidence, including one cut-marked ursid bone, illustrates the behavioural flexibility of Neanderthal groups at Fumane, who maintained a consistent focus on cervids while opportunistically exploiting other taxa when encountered. Collectively, the faunal evidence from Unit A9 highlights the planning depth, transport optimisation, and ecological sensitivity that characterised Neanderthal lifeways in northern Italy. These results reinforce a growing understanding across European Middle Palaeolithic contexts: Neanderthal foragers were sophisticated, behaviourally complex hunter-gatherers who structured their subsistence systems around high-utility carcass components, landscape constraints, and the energetic demands of Pleistocene environments.

Author Contributions

K.R.: Formal Analysis, Writing—Original Draft, Writing—Review and Editing, and Visualisation. N.N.: Formal Analysis, Validation, and Writing—Review and Editing. V.F. and M.D.L.: Formal Analysis. M.P.: Conceptualisation, Writing—Review and Editing, and Funding Acquisition. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding author(s).

Acknowledgments

Research at Fumane is coordinated by University of Ferrara in the framework of a project supported by the Ministry of Culture—Veneto Archaeological Superintendency, public institutions (Lessini Regional Natural Park, Fumane Municipality, MUSE-Museum of Science), and private associations and companies. This study was supported by University of Ferrara, FAR2023.

Conflicts of Interest

The authors declare no conflicts of interest.

References

  1. Romandini, M.; Nannini, N.; Tagliacozzo, A.; Peresani, M. The ungulate assemblage from layer A9 at Grotta di Fumane, Italy: A zooarchaeological contribution to the reconstruction of Neanderthal ecology. Quat. Int. 2014, 337, 11–27. [Google Scholar] [CrossRef]
  2. Delpiano, D.; Zupancich, A.; Peresani, M. Innovative Neanderthals: Results from an integrated analytical approach applied to backed stone tools. J. Archaeol. Sci. 2019, 110, 105011. [Google Scholar] [CrossRef]
  3. Delpiano, D.; Heasley, K.; Peresani, M. Assessing Neanderthal land use and lithic raw material management in Discoid technology. J. Anthropol. Sci. 2018, 96, 89–110. [Google Scholar] [CrossRef]
  4. Peresani, M.; Sauro, U. Palaeolithic Cave deposits and Karst evolution in the Venetian Pre-Alps. Geol. Field Trips Maps 2024, 16, 1–40. [Google Scholar] [CrossRef]
  5. Marcazzani, D.; Miller, C.E.; Ligouis, B.; Duches, R.; Conard, N.J.; Peresani, M. Middle and Upper Paleolithic occupations of Fumane Cave (Italy): A geoarchaeological investigation of the anthropogenic features. J. Anthropol. Sci. 2023, 101, 37–62. [Google Scholar]
  6. Peresani, M. Inspecting human evolution from a cave. Late Neanderthals and early sapiens at Grotta di Fumane: Present state and outlook. J. Anthropol. Sci. 2022, 100, 71–107. [Google Scholar]
  7. López-García, J.M.; dalla Valle, C.; Cremaschi, M.; Peresani, M. Reconstruction of the Neanderthal and Modern Human landscape and climate from the Fumane cave sequence (Verona, Italy) using small-mammal assemblages. Quat. Sci. Rev. 2015, 128, 1–13. [Google Scholar] [CrossRef]
  8. Kehl, M.; Marcazzan, D.; Miller, C.E.; Falcucci, A.; Duches, R.; Peresani, M. The Upper Sedimentary Sequence of Grotta di Fumane, Northern Italy: A Micromorphological Approach to Study Imprints of Human Occupation and Paleoclimate Change. Geoarchaeology 2025, 40, e70000. [Google Scholar] [CrossRef]
  9. Sánchez-Romero, L.; Govoni, M.; Marcazzan, D.; Delpiano, D.; Nannini, N.; Martellotta, E.F.; Duches, R.; Peresani, M. Intra-site Organization of the Repeated Neanderthal Occupation of Unit A9, Grotta di Fumane (Pre-Alps, Italy). J. Archaeol. Method Theory 2025, 32, 45. [Google Scholar] [CrossRef]
  10. France, D.L. Human and Nonhuman Bone Identification: A Color Atlas; CRC Press: New York, NY, USA, 2009. [Google Scholar]
  11. Davis, S.J.M. The Archaeology of Animals; Routledge Press: London, UK, 1987. [Google Scholar]
  12. Lyman, R.L. Vertebrate Taphonomy; Cambridge University Press: Cambridge, UK, 2014; pp. 1–524. [Google Scholar] [CrossRef]
  13. Broughton, J.M. Zooarchaeology. In International Encyclopedia of the Social & Behavioral Sciences, 2nd ed.; Elsevier: Amsterdam, The Netherlands, 2015; pp. 849–853. [Google Scholar]
  14. Reitz, E.J.; Wing, E.S. Cambridge Manual of Zooarchaology; Cambridge University Press: Cambridge, UK, 2008. [Google Scholar]
  15. Bunn, H.T.; Bartram, L.E.; Kroll, E.M. Variability in bone assemblage formation from Hadza hunting, scavenging, and carcass processing. J. Anthropol. Archaeol. 1988, 7, 412–457. [Google Scholar] [CrossRef]
  16. Aitken, R.J. Delayed implantation in roe deer (Capreolus capreolus). Reproduction 1974, 39, 225–233. [Google Scholar] [CrossRef]
  17. Mariezkurrena, K. Contribución al conocimiento del desarrollo de la dentición y el esqueleto postcraneal de Cervus elaphus. Munibe 1983, 35, 149–202. [Google Scholar]
  18. D’errico, F.; Vanhaeren, M. Criteria for Identifying Red Deer (Cervus elaphus) Age and Sex from Their Canines. Application to the Study of Upper Palaeolithic and Mesolithic Ornaments. J. Archaeol. Sci. 2002, 29, 211–232. [Google Scholar] [CrossRef]
  19. Hillson, S. Teeth; Cambridge University Press: Cambridge, UK, 2005. [Google Scholar]
  20. Silver, I. The Ageing of Domestic Animals in Science in Archaeology: A Survey of Progress and Research; Brothewell, D., Higgs, E., Clark, G., Eds.; Praeger Press: New York, NY, USA, 1969; pp. 250–268. [Google Scholar]
  21. Bunn, H.T.; Pickering, T.R. Methodological recommendations for ungulate mortality analyses in paleoanthropology. Quat. Res. 2010, 74, 388–394. [Google Scholar] [CrossRef]
  22. Habermehl, K.H. Die Altersbestimmung Bei Hausund Labortieren; Auflage: Berlin/Hamburg, Germany, 1975. [Google Scholar]
  23. Grayson, D.K. Quantitative Zooarchaeology: Topics in the Analysis of Archaeological Faunas; Academic Press: Orlando, FL, USA, 1984. [Google Scholar]
  24. Bökönyi, S. A New Method for the Determination of the Number of Individuals in Animal Bone Material. Am. J. Archaeol. 1970, 74, 291–292. [Google Scholar] [CrossRef]
  25. Behrensmeyer, A.K. Taphonomic and ecologic information from bone weathering. Paleobiology 1978, 4, 150–162. [Google Scholar] [CrossRef]
  26. Fernandez-Jalvo, Y.; Andrews, P. Atlas of Taphonomic Identifications; Springer: Dordrecht, The Netherlands, 2016. [Google Scholar]
  27. Brain, C.K. Bone weathering and the problem of bone pseudo-tools. S. Afr. J. Sci. 1967, 63, 97–99. [Google Scholar]
  28. Shipman, P. Life History of a Fossil: An Introduction to Taphonomy and Paleoecology; Harvard University Press: Cambridge, UK, 1981. [Google Scholar]
  29. Fisher, J.W. Bone surface modifications in zooarchaeology. J. Archaeol. Method Theory 1995, 2, 7–98. [Google Scholar] [CrossRef]
  30. Binford, L.R. Bones: Ancient Men and Modern Myth; Elsevier: Amsterdam, The Netherlands; Academic Press: Cambridge, MA, USA, 1981. [Google Scholar]
  31. Brain, C.K. The Hunters or the Hunted? An Introduction to African Cave Taphonomy; Chicago University Press: Chicago, IL, USA, 1981. [Google Scholar]
  32. Blumenschine, R.J.; Selvaggio, M.M. Percussion marks on bone surfaces as a new diagnostic of hominid behaviour. Nature 1988, 333, 763–765. [Google Scholar] [CrossRef]
  33. Capaldo, S.D.; Blumenschine, R.J. A Quantitative Diagnosis of Notches Made by Hammerstone Percussion and Carnivore Gnawing on Bovid Long Bones. Am. Antiq. 1994, 59, 724–748. [Google Scholar] [CrossRef]
  34. Blumenschine, R.J. Percussion marks, tooth marks, and experimental determinations of the timing of hominid and carnivore ace to long bones at FLK Zinjanthropus, Olduvai Gorge, Tanzania. J. Hum. Evol. 1995, 27, 197–213. [Google Scholar] [CrossRef]
  35. Domínguez-Rodrigo, M.; Piqueras, A. The use of tooth pits to identify carnivore taxa in tooth-marked archaeofaunas and their relevance to reconstruct hominid carcass processing behaviours. J. Archaeol. Sci. 2003, 30, 1385–1391. [Google Scholar] [CrossRef]
  36. Potts, R.; Shipman, P. Cutmarks made by stone tools on bones from Olduvai Gorge, Tanzania. Nature 1981, 291, 577–580. [Google Scholar] [CrossRef]
  37. Shipman, P.; Rose, J. Early hominid hunting, butchering, and carcass-processing behaviors: Approaches to the fossil record. J. Anthropol. Archaeol. 1983, 2, 57–98. [Google Scholar] [CrossRef]
  38. Lyman, R.L. Quantitative Paleozoology; Cambridge University Press: Cambridge, UK, 2008. [Google Scholar]
  39. Costamagno, S.; Soulier, M.-C.; Val, A.; Chong, S. The reference collection of cutmarks. Palethnologie 2019, 1, 1–93. [Google Scholar] [CrossRef]
  40. Blumenschine, R.J.; Selvaggio, M.M. On the marks of marrow bone processing by hammerstones hyaenas: Their anatomical patterning archaeological implications. In Cultural Beginnings: Approaches to Understanding Early Hominid Life-Ways in the African Savanna; Clark, J.D., Ed.; Dr Rudolf Habelt GMBH: Bonn, Germany, 1991; pp. 17–32. [Google Scholar]
  41. Blasco, R.; Rosell, J.; Domínguez-Rodrigo, M.; Lozano, S.; Pastó, I.; Riba, D.; Vaquero, M.; Peris, J.F.; Arsuaga, J.L.; de Castro, J.M.B.; et al. Learning by Heart: Cultural Patterns in the Faunal Processing Sequence during the Middle Pleistocene. PLoS ONE 2013, 8, e55863. [Google Scholar] [CrossRef]
  42. Vettese, D.; Daujeard, C.; Blasco, R.; Borel, A.; Caceres, I.; Moncel, M.H. Neandertal long bone breakage process: Standardized or random patterns? The example of Abri du Maras (Southeastern France, MIS 3). J. Archaeol. Sci. Rep. 2017, 13, 151–163. [Google Scholar] [CrossRef]
  43. Villa, P.; Mahieu, E. Breakage patterns of human long bones. J. Hum. Evol. 1991, 21, 27–48. [Google Scholar] [CrossRef]
  44. Outram, A.K. A New Approach to Identifying Bone Marrow and Grease Exploitation: Why the “Indeterminate” Fragments should not be Ignored. J. Archaeol. Sci. 2001, 28, 401–410. [Google Scholar] [CrossRef]
  45. Grunwald, A.M. Analysis of fracture patterns from experimentally marrow-cracked frozen and thawed cattle bones. J. Archaeol. Sci. Rep. 2016, 8, 356–365. [Google Scholar] [CrossRef]
  46. Coil, R.; Tappen, M.; Yezzi-Woodley, K. New analytical methods for comparing bone fracture angles: A controlled study of hammerstone and hyena (Crocuta crocuta) long bone breakage. Archaeometry 2017, 59, 900–917. [Google Scholar] [CrossRef]
  47. Stiner, M.C.; Kuhn, S.L.; Weiner, S.; Bar-Yosef, O. Differential Burning, Recrystallization, and Fragmentation of Archaeological Bone. J. Archaeol. Sci. 1995, 22, 223–237. [Google Scholar] [CrossRef]
  48. Miller, F.L. Caribou. In Wild Mammals of North America: Biology, Management, and Conservation; Feldhamer, G.A., Thompson, B.C., Chapman, J.A., Eds.; John Hopkins University Press: Baltimore, MD, USA, 2003; pp. 965–997. [Google Scholar]
  49. Mysterud, A. Diet overlap among ruminants in Fennoscandia. Oecologia 2000, 124, 130–137. [Google Scholar] [CrossRef]
  50. Jackson, J.E. Feeding habits of deer. Mammal Rev. 1974, 4, 93–101. [Google Scholar] [CrossRef]
  51. Alekseĭ, D.; Hewison, A.M. Behavioural Ecology of Siberian and European Roe Deer; Chapman & Hall: London, UK, 1996. [Google Scholar]
  52. van Beeck Calkoen, S.T.S.; Leigh-Moy, K.; Cromsigt, J.P.G.M.; Spong, G.; Lebeau, L.C.; Heurich, M. The blame game: Using eDNA to identify species-specific tree browsing by red deer (Cervus elaphus) and roe deer (Capreolus capreolus) in a temperate forest. For. Ecol. Manag. 2019, 451, 117483. [Google Scholar] [CrossRef]
  53. Yravedra, J.; Cobo-Sánchez, L. Neanderthal exploitation of ibex and chamois in southwestern Europe. J. Hum. Evol. 2015, 78, 12–32. [Google Scholar] [CrossRef] [PubMed]
  54. Carvalho, M.; Haws, J.A.; Jones, E.L. Late Neanderthal subsistence foraging mobility at Lapa do Picareiro: A zooarchaeological taphonomic analysis of Level JJ. Front. Environ. Archaeol. 2025, 4, 1–21. [Google Scholar] [CrossRef]
  55. Rendu, W.; Costamagno, S.; Meignen, L.; Soulier, M.-C. Monospecific faunal spectra in Mousterian contexts: Implications for social behavior. Quat. Int. 2012, 247, 50–58. [Google Scholar] [CrossRef]
  56. Bal-García, H.; Rey-Rodríguez, I.; de Lombera-Hermida, A.; Díaz-Rodríguez, M.; Fernández-Rodríguez, C.; Rodríguez-Álvarez, X.P.; Valcarce, R.F. Landscape and subsistence in NW Iberia during the Middle Palaeolithic (MIS3): Faunal analysis of Cova Eirós (Triacastela, Galicia, Spain). J. Archaeol. Sci. Rep. 2025, 64, 105149. [Google Scholar] [CrossRef]
  57. Yravedra, J.; Estaca-Gómez, V.; Grandal-d’Anglade, A.; Pinto-Llona, A.C. Neanderthal subsistence strategies: New evidence from the Mousterian Level XV of the Sopeña rock shelter (Asturias, northern Spain). Archaeol. Anthropol. Sci. 2024, 16, 6. [Google Scholar] [CrossRef]
  58. Grimstead, D.N. Ethnographic and Modeled Costs of Long-Distance, Big-Game Hunting. Am. Antiq. 2010, 75, 61–80. [Google Scholar] [CrossRef]
  59. Grimstead, D.N. Prestige and Prejudice: The Role of Long Distance Big Game Hunting as an Optimal Foraging Decision. Am. Antiq. 2012, 77, 168–178. [Google Scholar] [CrossRef]
  60. Broughton, J.M.; Cannon, M.D.; Bayham, F.E.; Byers, D.A. Prey Body Size and Ranking in Zooarchaeology: Theory, Empirical Evidence, and Applications from the Northern Great Basin. Am. Antiq. 2011, 76, 403–428. [Google Scholar] [CrossRef]
  61. Bird, D.W.; Bird, R.B.; Codding, B.F. In Pursuit of Mobile Prey: Martu Hunting Strategies and Archaeofaunal Interpretation. Am. Antiq. 2009, 74, 3–29. [Google Scholar] [CrossRef][Green Version]
  62. Wilson, D.E.; Mittermeier, R.A. (Eds.) Handbook of the Mammals of the World, Volume 1: Carnivores; Lynx Edicions: Barcelona, Spain, 2009. [Google Scholar]
  63. López-González, F.; Grandal-d’Anglade, A.; Vidal-Romaní, J.R. Deciphering bone depositional sequences in caves through the study of manganese coatings. J. Archaeol. Sci. 2006, 33, 707–717. [Google Scholar] [CrossRef]
  64. Pineda, A.; Saladié, P. Beyond the Problem of Bone Surface Preservation in Taphonomic Studies of Early and Middle Pleistocene Open-Air Sites. J. Archaeol. Method Theory 2022, 29, 1090–1130. [Google Scholar] [CrossRef]
  65. Schiffer, M.B. Toward the Identification of Formation Processes. Am. Antiq. 1983, 48, 675–706. [Google Scholar] [CrossRef]
  66. Morin, E. Taphonomy. In Reassessing Paleolithic Subsistence The Neandertal and Modern Human Foragers of Saint-Césaire; Cambridge University Press: Cambridge, UK, 2012; pp. 93–131. [Google Scholar]
  67. Macho-Callejo, A.; García-Morato, S.; Gutiérrez, A.; Marin-Monfort, D.; Fernández-Jalvo, Y. Put down roots and find the plant!: Preliminary results of root etching and its implications. Hist. Biol. 2024, 36, 2502–2510. [Google Scholar] [CrossRef]
  68. Olsen, S.L.; Shipman, P. Surface modification on bone: Trampling versus butchery. J. Archaeol. Sci. 1988, 15, 535–553. [Google Scholar] [CrossRef]
  69. Moclán, A.; Domínguez-Rodrigo, M. An experimental study of the patterned nature of anthropogenic bone breakage and its impact on bone surface modification frequencies. J. Archaeol. Sci. 2018, 96, 1–13. [Google Scholar] [CrossRef]
  70. Thun Hohenstein, U.; Caffarelli, L.; Arnetta, G.; Rivals, F.; Pozzobon, P.; Gialanella, S.; Delpiano, D.; Peresani, M. Taphonomy of the fauna and chert assemblages from the Middle Palaeolithic site of Vajo Salsone, Eastern Italian Alps. Quat. Sci. Adv. 2024, 14, 100183. [Google Scholar] [CrossRef]
  71. Modolo, M.; Delpiano, D.; Martellotta, E.F.; Peresani, M. Bone Refits and Implications for the Reconstruction of a Late Middle Palaeolithic Context: Unit A9 of Fumane Cave. J. Paleolit. Archaeol. 2024, 8, 1. [Google Scholar] [CrossRef]
  72. Thun-Hohenstein, U.; Parere, V.; Sala, B.; Giunti, P.; Longo, L. Large mammals from Mezzena rockshelter: New biocronological and palaeoecological hypotheses and preliminary data on subsistence strategies. Quat. Int. 2012, 259, 1–6. [Google Scholar]
  73. Fiore, I.; Gala, M.; Romandini, M.; Cocca, E.; Tagliacozzo, A.; Peresani, M. From feathers to food: Reconstructing the complete exploitation of avifaunal resources by Neanderthals at Fumane cave, unit A9. Quat. Int. 2016, 421, 134–153. [Google Scholar] [CrossRef]
  74. Romandini, M.; Hohenstein, U.T.; Fiore, I.; Tagliacozzo, A.; Perez, A.; Lubrano, V.; Terlato, G.; Peresani, M. Late neandertals and the exploitation of small mammals in northern Italy: Fortuity, necessity or hunting variability? Quaternaire 2018, 29, 61–67. [Google Scholar] [CrossRef]
  75. Terlato, G.; Livraghi, A.; Romandini, M.; Peresani, M. Large bovids on the Neanderthal menu: Exploitation of Bison priscus and Bos primigenius in northeastern Italy. J. Archaeol. Sci. Rep. 2019, 25, 129–143. [Google Scholar] [CrossRef]
  76. Schoville, B.J.; Otárola-Castillo, E. A model of hunter-gatherer skeletal element transport: The effect of prey body size, carriers, and distance. J. Hum. Evol. 2014, 73, 1–14. [Google Scholar] [CrossRef]
  77. Faith, J.T.; Gordon, A.D. Skeletal element abundances in archaeofaunal assemblages: Economic utility, sample size, and assessment of carcass transport strategies. J. Archaeol. Sci. 2007, 34, 872–882. [Google Scholar] [CrossRef]
  78. Binford, L.R. Nunamiut Ethnoarchaeology; Academic Press: New York, NY, USA, 1978. [Google Scholar]
  79. Bunn, H.T. Patterns of skeletal representation and hominid subsistence activities at Olduvai Gorge, Tanzania, and Koobi Fora, Kenya. J. Hum. Evol. 1986, 15, 673–690. [Google Scholar] [CrossRef]
  80. Bartram, L.E. Perspectives on skeletal part profiles and utility curves from eastern Kalahari ethnoarchaeology. In From Bones to Behavior: Ethnoarchaeological and Experimental Contributions to the Interpretation of Faunal Remains; Hudson, J., Ed.; The Center for Archaeological Investigations at Southern Illinois University: Carbondale, IL, USA, 1993; pp. 115–137. [Google Scholar]
  81. Lam, Y.M.; Chen, X.; Pearson, O.M. Intertaxonomic Variability in Patterns of Bone Density and the Differential Representation of Bovid, Cervid, and Equid Elements in the Archaeological Record. Am. Antiq. 1999, 64, 343–362. [Google Scholar] [CrossRef]
  82. Lyman, R.L. Anatomical considerations of utility curves in Zooarchaeology. J. Archaeol. Sci. 1992, 19, 7–22. [Google Scholar] [CrossRef]
  83. Binford, L.R. In Pursuit of the Past: Decoding the Archaeological Record; University of California Press: Los Angeles, CA, USA, 1983. [Google Scholar]
  84. Chang, C.-Y.; Ke, D.-S.; Chen, J.-Y. Essential fatty acids and human brain. Acta Neurol. Taiwanica 2009, 18, 231–241. [Google Scholar]
  85. Agam, A.; Barkai, R. Not the brain alone: The nutritional potential of elephant heads in Paleolithic sites. Quat. Int. 2016, 406, 218–226. [Google Scholar] [CrossRef]
  86. Rougier, H.; Crevecoeur, I.; Beauval, C.; Posth, C.; Flas, D.; Wißing, C.; Furtwängler, A.; Germonpré, M.; Gómez-Olivencia, A.; Semal, P.; et al. Neandertal cannibalism and Neandertal bones used as tools in Northern Europe. Sci. Rep. 2016, 6, 29005. [Google Scholar] [CrossRef]
  87. Domínguez-Rodrigo, M.; Pickering, T.R.; Semaw, S.; Rogers, M.J. Cutmarked bones from Pliocene archaeological sites at Gona, Afar, Ethiopia: Implications for the function of the world’s oldest stone tools. J. Hum. Evol. 2005, 48, 109–121. [Google Scholar] [CrossRef]
  88. Blasco, R.; Fernández Peris, J. A uniquely broad spectrum diet during the Middle Pleistocene at Bolomor Cave (Valencia, Spain). Quat. Int. 2012, 252, 16–31. [Google Scholar] [CrossRef]
  89. Marín, J.; Daujeard, C.; Saladié, P.; Rodríguez-Hidalgo, A.; Vettese, D.; Rivals, F.; Boulbes, N.; Crégut-Bonnoure, E.; Lateur, N.; Gallotti, R.; et al. Neanderthal faunal exploitation and settlement dynamics at the Abri du Maras, level 5 (south-eastern France). Quat. Sci. Rev. 2020, 243, 106472. [Google Scholar] [CrossRef]
  90. Chase, P.G. The Hunters of Combe-Grenal: Approaches to Middle Paleolithic Subsistence in Europe. In British Archaeological Reports; BAR International Series S286; BAR Publishing: Oxford, UK, 1986. [Google Scholar]
  91. Berlioz, E.; Capdepon, E.; Discamps, E. A long-term perspective on Neanderthal environment and subsistence: Insights from the dental microwear texture analysis of hunted ungulates at Combe-Grenal (Dordogne, France). PLoS ONE 2023, 18, e0278395. [Google Scholar] [CrossRef]
  92. Romandini, M.; Terlato, G.; Nannini, N.; Tagliacozzo, A.; Benazzi, S.; Peresani, M. Bears and humans, a Neanderthal tale. Reconstructing uncommon behaviors from zooarchaeological evidence in southern Europe. J. Archaeol. Sci. 2018, 90, 71–91. [Google Scholar] [CrossRef]
  93. Livraghi, A.; Fanfarillo, G.; Colle, M.D.; Romandini, M.; Peresani, M. Neanderthal ecology and the exploitation of cervids and bovids at the onset of MIS4: A study on De Nadale cave, Italy. Quat. Int. 2021, 586, 24–41. [Google Scholar] [CrossRef]
  94. Romandini, M.; Silvestrini, S.; Real, C.; Lugli, F.; Tassoni, L.; Carrera, L.; Badino, F.; Bortolini, E.; Marciani, G.; Delpiano, D.; et al. Late Neanderthal “menu” from northern to southern Italy: Freshwater and terrestrial animal resources. Quat. Sci. Rev. 2023, 315, 108233. [Google Scholar] [CrossRef]
  95. Gaudzinski, S. A matter of high resolution? The Eemian Interglacial (OIS 5e) in north-central Europe and Middle Palaeolithic subsistence. Int. J. Osteoarchaeol. 2004, 14, 201–211. [Google Scholar] [CrossRef]
  96. Adler, D.S.; Bar-Oz, G.; Belfer-Cohen, A.; Bar-Yosef, O. Ahead of the Game. Curr. Anthropol. 2006, 47, 89–118. [Google Scholar] [CrossRef]
  97. Stiner, M.C.; Munro, N.D.; Surovell, T.A. The Tortoise and the Hare: Small-Game Use, the Broad-Spectrum Revolution, and Paleolithic Demography. Curr. Anthropol. 2000, 41, 39–79. [Google Scholar] [CrossRef]
  98. Uzunidis, A.; Blasco, R.; Brugal, J.-P.; Fourcade, T.; Ochando, J.; Rosell, J.; Roussel, A.; Rufà, A.; Sánchez Goñi, M.F.; Texier, P.-J.; et al. Neanderthal hunting grounds: The case of Teixoneres Cave (Spain) and Pié Lombard rockshelter (France). J. Archaeol. Sci. 2024, 168, 106007. [Google Scholar] [CrossRef]
  99. Daujeard, C.; Vettese, D.; Britton, K.; Béarez, P.; Boulbes, N.; Crégut-Bonnoure, E.; Desclaux, E.; Lateur, N.; Pike-Tay, A.; Rivals, F.; et al. Neanderthal selective hunting of reindeer? The case study of Abri du Maras (south-eastern France). Archaeol. Anthropol. Sci. 2019, 11, 985–1011. [Google Scholar] [CrossRef]
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Figure 1. (A) location of Fumane Cave in the north-east of Italy (red frame in the map inlet) and showing major Middle Palaeolithic cave sites in the Lessini Mountains and Berici Hill; (B) plan of the cave excavation area, with the innermost grid squares sampled in this study indicated (modified by D. Delpiano).
Figure 1. (A) location of Fumane Cave in the north-east of Italy (red frame in the map inlet) and showing major Middle Palaeolithic cave sites in the Lessini Mountains and Berici Hill; (B) plan of the cave excavation area, with the innermost grid squares sampled in this study indicated (modified by D. Delpiano).
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Figure 2. Natural modifications observed in the assemblage.
Figure 2. Natural modifications observed in the assemblage.
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Figure 3. Ungulate distribution (NISP) across the faunal assemblage.
Figure 3. Ungulate distribution (NISP) across the faunal assemblage.
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Figure 4. Representation of the frequency of anatomical elements for the main taxa of ungulates.
Figure 4. Representation of the frequency of anatomical elements for the main taxa of ungulates.
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Figure 5. Cut marks on ungulate diaphysis (A9-RF 264) (left) and cut marks (skinning action) on Cervus elaphus diaphysis (A9-RF 155) (right).
Figure 5. Cut marks on ungulate diaphysis (A9-RF 264) (left) and cut marks (skinning action) on Cervus elaphus diaphysis (A9-RF 155) (right).
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Figure 6. Percussion marks and cut marks on caprine tibia (RF 046) (left) and retouching marks on Cervus elaphus tibia (RF 002) (right) from Unit A9.
Figure 6. Percussion marks and cut marks on caprine tibia (RF 046) (left) and retouching marks on Cervus elaphus tibia (RF 002) (right) from Unit A9.
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Figure 7. Carcass representation profile of ungulates, Fumane Unit A9.
Figure 7. Carcass representation profile of ungulates, Fumane Unit A9.
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Table 1. Taxonomical determinations of the assemblage followed by NISP and MNI.
Table 1. Taxonomical determinations of the assemblage followed by NISP and MNI.
TaxaNISPNISP%MNI
Rodentia indet.40.91
Total Rodentia40.91
Aves10.21
Total Aves10.21
Vulpes vulpes10.21
Ursus arctos30.73
Ursus spelaeus20.52
Ursus sp.30.73
Meles meles20.51
Carnivora indet10.2
Total Carnivora122.810
Megaloceros giganteus10.21
Alces alces20.51
Cervus elaphus9221.78
Capreolus capreolus307.13
Large Cervidae30.72
Cervidae indet.122.85
Caprine61.41
Capra ibex81.93
Rupicapra rupicapra184.22
Ungulata indet.23555.4
Total Ungulates4079626
Total Determined42424.438
Undetermined Specimens
Small-sized mammals10.1
Medium-sized mammals181.0
Medium–big-sized mammals271.6
Big-sized mammals80.5
Undetermined diaphysis126072.5
Total Undetermined131475.6
Total NISPs1738100.00
Table 2. Ungulates with butchering marks, thermal alterations, and carnivore tooth marks. CM: cut marks; PM: percussion marks; CM + PM: cut marks + percussion marks; Tot. BM: total butchering marks; % BM: % butchering marks; R: retoucher; B: burned-black/brown bones; C: calcined-grey/white bones; TM: carnivore tooth marks.
Table 2. Ungulates with butchering marks, thermal alterations, and carnivore tooth marks. CM: cut marks; PM: percussion marks; CM + PM: cut marks + percussion marks; Tot. BM: total butchering marks; % BM: % butchering marks; R: retoucher; B: burned-black/brown bones; C: calcined-grey/white bones; TM: carnivore tooth marks.
TaxaNISPCMPMCM + PMTot. BM% BMRBCTM
Megaloceros giganteus1
Alces alces21 150.00 1
Cervus elaphus9228413335.871337
Capreolus capreolus309111136.67 11
Large Cervidae3
Cervidae indet.121 18.33
Caprine6211466.67
Capra ibex821 337.50
Rupicapra rupicapra18311527.7812
Ungulata indet.23565226929.362685
Total40711110612731.204131212
Table 3. Number of anatomical elements of Cervus elaphus with anthropic modifications and carnivore tooth marks. CM: cut marks; PM: percussion marks; CM + PM: cut marks and percussion marks; CM + IF: cut marks + impact flakes; Tot. BM: total butchering marks; % BM: % butchering marks; R: retoucher; B: burned-black/brown bones; C: calcined-grey/white bones; TM: carnivore tooth marks.
Table 3. Number of anatomical elements of Cervus elaphus with anthropic modifications and carnivore tooth marks. CM: cut marks; PM: percussion marks; CM + PM: cut marks and percussion marks; CM + IF: cut marks + impact flakes; Tot. BM: total butchering marks; % BM: % butchering marks; R: retoucher; B: burned-black/brown bones; C: calcined-grey/white bones; TM: carnivore tooth marks.
Anatomical ElementNISPCMPMCM + PMTot. BM% BMRBCTM
Cranium
Hemimandible51 120.0 1 1
Tooth4
Hyoid
Total cranium91 111.1
Atlas axis
Vertebra
Rib
Total axial skeleton
Scapula
Humerus62 233.3 11
Radius53 360.0
Ulna
Metacarpal144 428.6 1 2
Total forelimb259 936.0
Coxal1
Femur12331758.3 1
Tibia134 430.81 2
Metatarsal126 650.0 111
Total hindlimb3813311744.7
Metapodials621 350.0
First phal. 72 228.6
Second phal.
Third phal. 2
First phal. rudim.
Second phal. rudim. 1
Third phal. rudim. 1
Sesamoid21 150.0
Diaphysis1
Total indet. Limb2051 630.0
Total9228413335.91337
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Rodrigo, K.; Nannini, N.; Facincani, V.; De Lorenzi, M.; Peresani, M. From Prey to Pattern: Integrating Faunal and Behavioural Evidence of Neanderthal Subsistence at Fumane Cave (Unit A9), Northern Italy. Quaternary 2026, 9, 14. https://doi.org/10.3390/quat9010014

AMA Style

Rodrigo K, Nannini N, Facincani V, De Lorenzi M, Peresani M. From Prey to Pattern: Integrating Faunal and Behavioural Evidence of Neanderthal Subsistence at Fumane Cave (Unit A9), Northern Italy. Quaternary. 2026; 9(1):14. https://doi.org/10.3390/quat9010014

Chicago/Turabian Style

Rodrigo, Kalangi, Nicola Nannini, Vittorio Facincani, Matteo De Lorenzi, and Marco Peresani. 2026. "From Prey to Pattern: Integrating Faunal and Behavioural Evidence of Neanderthal Subsistence at Fumane Cave (Unit A9), Northern Italy" Quaternary 9, no. 1: 14. https://doi.org/10.3390/quat9010014

APA Style

Rodrigo, K., Nannini, N., Facincani, V., De Lorenzi, M., & Peresani, M. (2026). From Prey to Pattern: Integrating Faunal and Behavioural Evidence of Neanderthal Subsistence at Fumane Cave (Unit A9), Northern Italy. Quaternary, 9(1), 14. https://doi.org/10.3390/quat9010014

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