The fossil traces of Stone Age humans and other animals in the Grotta della Bàsura cave system in Italy have been studied since the 1950s. Italian archaeologist Virginia Chiappella published the first studies; she documented bones from an extinct cave bear, human and animal footprints, charcoal from torches, finger marks, and lumps of clay stuck on the walls. Since then, many more archeologists and anthropologists have studied the cave and its fossils. Yet there are still lessons to be learned from this prehistoric site.

Now, Romano et al. have combined a number of different approaches and used some of the latest technology and cutting-edge software to analyze 180 footprints and other tracks found in the cave. These trace fossils date to about 14,000 years ago, and the analysis revealed that they were left by a group of Stone Age humans who descended at least 400 meters into the cave. The group consisted of two adults, an adolescent and two children of about three and six years old. At one point they had to crawl through a low tunnel – something that has not previously been documented in the fossil record. The group were all barefoot, had no clothing on their arms and legs and used wooden torches to light the way.

Together, these findings suggest that young children were active group members during the late Stone Age, even when carrying out apparently dangerous activities. Romano et al. now hope that their multidisciplinary approach may help other scientists looking to understand how humans behaved elsewhere in the world at various points in history.

Discovered in 1950, the hypogeal part of the ‘Grotta della Bàsura’ is a large and deep cave that has produced some of the most important Italian Palaeolithic discoveries of the twentieth century (Chiappella, 1952; Tongiorgi and Lamboglia, 1954; Blanc, 1960; Lamboglia, 1960; Giacobini, 2008) consisting of traces of human activity, especially footprints, dating back to 14 ky cal. BP. Up till 1890 only the atrial part of the cavity was known where Neolithic and late Roman archaeological finds had been unearthed (Maineri, 1985). The cave opening is situated at 186 m a.s.l. about 1 km north of Toirano (Savona, Italy - 436253.433 E; 4887689.739 N), and extends 890 m into Mount S. Pietro with an elevation difference of +20/−22 m relative to that of the entrance (Figure 1).

Figure 1

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The inner rooms became accessible in 1950, after the rupture of a stalagmite column, placed a few dozen meters from the entrance, that prevented any access to the cave (Tongiorgi and Lamboglia, 1954; Blanc, 1960; Lamboglia, 1960). The exceptional prehistoric and paleontological value of the cave was firstly recognized by Virginia Chiappella (Chiappella, 1952) who was the first scholar to visit the site soon after its discovery. Chiappella identified several bones of Ursus spelaeus and traces both of animal and human frequentation (footprints, charcoals, digital tracks, lumps of clay adhering to the walls) in different areas of the cave within approximately 350 m of the entrance. Regrettably, destruction of most of the ichnological record occurred as a result of uncontrolled cave visits by local villagers and tourists lured to the remarkable discovery as a result of media reports (Blanc et al., 1960; De Lumley and Giacobini, 1985).

The first study of human footprints from the Cave of Bàsura was conducted by Pales (1960), based on original images and 13 plaster casts of the best-preserved specimens from various sectors of the cave. Analysis of the bone architecture of the footprint makers and the apparent relationship with remains of Ursus spelaeus, led the author to consider that the footprints were made by ‘Neanderthal-type’ producers. Subsequent analysis (de Lumley and Vicino, 1984), accompanied by radiometric dating, placed the prehistoric visitation of the Bàsura in the Upper Palaeolithic, between 12,000 and 14,000 years uncal BP (de Lumley and Vicino, 1984; De Lumley and Giacobini, 1985). Dating of charred wood fragments found on the trampled surface provided a more precise age for cave visitation at 12,340 years ± 160 years BP (Molleson et al., 1972; Molleson, 1985).

Based on the ichnological study of Pales, Blanc proposed that the inner room called ‘Sala dei Misteri’ was reached by multiple individuals, including a juvenile. Blanc (1960) described, from the same room, a group of seven human footprints identified by heel tracks positioned only a few centimeters from the main wall of the chamber to which numerous lumps of clay were attached. The close association of the footprints with lumps of clay resulted in it being considered the result of initiation rites involving young hunters. This hypothesis was supported by the presence of a stalagmite concretion (defined by Blanc as ‘acephalous sphinx’ or ‘zoomorphic stalagmite’) placed against the terminal wall of the ‘Sala dei Misteri’ (Mysteries’ Hall) chamber on which several sinuous furrows had been made intentionally by several individuals using their fingers. This interpretation is currently under review by two of the present authors (ES, MZ). The aim of this work is to review and improve understanding of the Grotta della Bàsura’s human ichnology, soil micromorphology, sedimentology and radiocarbon chronology. A preliminary report from this study focused on human footprints preserved in the innermost chamber of the cave (Citton et al., 2017).

In the present paper, we expanded our study to analyze and interpret all human traces (footprints and handprints as well as other traces) from the ‘Grotta della Bàsura’, providing new insight on the behavior, identity of the members, their exploratory techniques, and the social structure of an Upper Palaeolithic group.

Data

A total of 180 footprints and traces sensu lato were recorded and studied (Supplementary file 1). In addition to footprints (Figure 2), among the traces are digit and handprints on the clay-rich floor, and smears from hands dirtied with charcoal on the side walls of the cave (Figure 3) (Giannotti, 2008). A paleo-archaeological excavation was performed in 2016 in the ‘Sala dei Misteri’. This survey highlighted the total absence of archaeological material but led to the recovery, on the trampled palaeosurface, of numerous charcoal remains with bundles of Pinus t. sylvestris/mugo originally used to illuminate the cave. New radiometric dating on these charcoal samples constrains the exploration of the cave to the late Upper Palaeolithic, between 12,310 ± 60 and 12,370 ± 60 BP, that is from about 14,700 to 14,000 cal BP (Table 1).

Figure 2

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Figure 3

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Table 1

Sample name	Provenance	Dated material	Description	Lab. code	F14C ± 1σ	14C Age (yr BP) ±1σ	Calibrated* dating result (95.4 probability)	%C	%N	δ13C (in ‰;±1σ)	δ15N (in ‰;±1σ)	C/N ratio
Bàsura SM B5 17B	‘sala dei misteri’, square B5, Unit 1	Charcoal (AAA)	Pinus t. sylvestris/mugo	GrA-69598	0.2160 ± 0.0016	12310 ± 60	12720–12110 cal BC	71.4	-	‐25.24 ± 0.14	-	-
Bàsura SM D6 33B	‘Sala dei misteri’, square D6, Unit 1	charcoal (AAA)	Pinus t. sylvestris/mugo	GrA-69597	0.2145 ± 0.0016	12370 ± 60	12830–12165 cal BC	63.3	-	‐26.37 ± 0.14	-	-

Digit and hand traces are preserved in several sectors of the cave (Figure 3). Most are unintentional traces related to cave exploration activities (Figure 3, C0, C26b, C72). Others, especially in the inner chamber (‘Sala dei Misteri’) which are still being studied, are very probably related to social or symbolic activities and can be instead considered intentional (Figure 3, SM55, SM56). Moreover, different sized bear and Canidae incertae sedis footprints are ubiquitously present (Figure 4), and are often associated with human prints. The available data regarding the footprints attributed to ‘canids’ suggest a very reduced number of individuals and a close association with the human prints. Should ongoing studies (Avanzini et al in prep.) confirm that the ichnnological association of Bàsura could prove crucial to shed new light on dog domestication in the Upper Paleolithic (Morey and Jeger, 2015; Perri, 2016; Lupo, 2017; Janssens et al., 2018).

Figure 4

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Ursus sp. hibernation areas are still recognizable with well-preserved nests of both cubs and adult bears.

Geometric morphometry performed on human footprints highlighted five main morphotypes (hereafter Morphs.) indicating a possible minimum number of five individuals entering the cave (Figure 5). This number is confirmed by the construction of morphological groups (Table 2) reconstructed through the overlapping of footprints that show a variability of less than 2% of the main parameters.

Figure 5 with 1 supplement see all

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Morphs. 1 and 2 can be easily distinguished on the basis of the absolute footprint size. Morph. 1 includes footprints with a length of 13.55 ± 0.49 cm showing characters indicative of an early ontogenetic stage of the producer, such as digit traces and the heel area proportionally wider than those of the longer tracks. Morph. 2, with a length of 17 cm, is distinguished from Morph. one on the basis of a more pronounced plantar arch. Morph. 3 comprises footprints 20.83 ± 0.51 cm in length (Figure 5 and Table 3). The plantar area is characterized by a very pronounced medial embayment. This corresponds with a strongly convex external margin, as is shared together with a strong adduction of digit I trace, an overall larger divarication of digit traces and a consistent separation between adjacent digits II-III and IV-V.

Morph. 4 (Figures 5 and 6) is represented by larger footprints (22.80 ± 0,42 cm in overall length) with roughly straight medial and lateral margins and a medial embayment less marked than that of Morph. 3. Digit tip traces are strongly aligned and oriented forward. Morph. 5 (Figures 5 and 7) includes footprints of 25.73 ± 0.45 cm in overall length and slightly concave margins, with a variably pronounced plantar embayment. The footprints of Morph. 5 are generally more robust and stockier than those of Morphs. 3 and 4, sharing with Morph. 4 the straight, forwardly oriented digit tips, and with Morph. 3 an adducted digit I trace.

Figure 6

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Figure 7

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Plantigrade tracks enabled us to estimate stature, weight and ontogenetic stage of the producers based on biometric measurements (Table 3) and the adopted formulas (see Methods). An estimation of the gender for Morph. 5 was also attempted (see Methods).

The group of human track producers entering the cave comprised: a three-year-old child about 88 cm tall (Morph. 1); a child at least 6 years old and about 110 cm tall (Morph. 2); a pre-adolescent, between 8 and 11 years old, about 135 cm tall (Morph. 3); a sub-adult to adult about 148 cm tall (Morph. 4); and an adult about 167 cm tall (Morph. 5). Estimate of the stature for the Morph. 5 is further supported by the length of the tibia derived from the available kneeling traces (see Methods). Our results concerning Morphs. 4 and 5, which are referred to adult individuals, are in agreement with the average stature of European Upper Palaeolithic people (162.4 ± 4.6 cm for males and 153.9 ± 4.3 cm for females) (Villotte et al., 2017).

Body mass estimates derived from footprint parameters suggests slender and muscular body proportions for all the trackmakers. Arch angle and footprint morphology suggests a male as the probable trackmaker of the largest footprint group. Differently, the gender result difficult to infer for morphotypes 1, 2, 3, 4, although Morph. 4 can be referred most probably to a female.

Digitigrade and semi-plantigrade footprints provide information on pedal postures and the behavior of the producers passing through different sub-environments of the cave. Both these footprint types were, in most cases, traced back to the same type of producer by comparison with complete footprints indicating complete foot support during locomotion. Some semi-plantigrade footprints (e.g. Figure 8, C44, C44b) show a strongly adducted trace of digit I and an apparent alignment with the other digits, probably because of the intra-rotation movements of the distal portion of the foot during the thrust phase. Footprints included in the Morph. 3 show a peculiar pedal morphology. While the resulting morphology of the digit I trace is explained by walking on a waterlogged substrate, the separation between digit pairs II-III and IV-V suggest an inherited familiar trait or a pathological condition of the producer’s feet. The producer was not incapacitated, showing the greatest mobility in the hypogeal environment.

Figure 8

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In the lower corridor (Figure 9), a few of these footprints are associated with elongated traces imprinted by the producers’ knees resting on the substrate. Successions of kneeling traces can be clearly recognized for Morphs. 3, 4, and 5. Based on the overall size of the metatarsal and knee couples aligned on the substrate, crawling in a totally unknown environment is inferred for the whole group. These knee imprints (e.g. Figure 8, C42) show the muscle structure of the knee joint and adjacent regions. The patella, the patellar ligament (tendon), the tibial tuberosity, the fibular head, the basis of the vastus medial and the iliotibial band are recognizable and allow us to infer the body structure of the trackmakers.

Figure 9

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Successions of kneeling traces allowing one to infer a crawling locomotion for the trackmakers have never been reported before. Isolated kneeling traces (i.e. a footprint followed by a knee imprint of the same leg) have only previously been reported from the ‘Galerie Wahl’ at Fontanet cave and from the ‘Salle des Talons’ at Tuc d’Audoubert cave, France (Pastoors et al., 2015) but are not sufficient to infer crawling. In addition, the anatomic details clearly recognizable in the crawling traces from ‘Grotta della Bàsura’ enabled us to hypothesize that no clothing was interposed between the limb and the trampled sediments.

The integration of all available ichnological evidence with data on the complex morphology of the cave enabled us to reconstruct a detailed hypothesis for the events which took place by Paleolithic peoples while exploring the cave about 14,000 BP.

The short radiometric interval, documented by both previous radiocarbon dates and those derived from the latest research presented here (Table 1), together with interferences relating to the interrelations of the different footprints, suggest that all individuals entered the cave at the same time. In particular, the group of footprints C34, C35b, C36, C35 in the terminal part of the ‘Corridoio delle impronte’ provides relative timing of the impressions: C35 (Morph. 4) is superimposed on C36 (Morph. 3) and on C34 and C35b (both Morph. 5) (Figure 9, d3). In this specific case, the producer of Morph. 4 passed after the producers of Morphs 3 and 5, respectively. In the entrance of the same corridor, the timing is reversed: footprint C63 (Morph. 4) is superimposed by C64 (Morph. 3) proving that Morphs. 3 and 4 were made at the same time (Figure 10).

Figure 10

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It is also difficult to assume that Morphs. 1 and 2 may have entered the deepest part of the cave without the presence of some older individual: if the older individuals who were leading the exploration were either Morphs. 3 or 4 the group consequently increases to include at least four individuals. No relationship can be established for the larger individual; however, as reported below, the fact that the footprints of this individual are regular and were closely followed on the same path by all the other individuals, suggests that the larger one was likely the leader of the synchronous exploration. Thus, all the available lines of evidence, particularly the complex interrelationship of the studied footprints, strongly suggest that a single exploration event by a heterogeneous group of five individuals is the most parsimonious and best supported hypothesis.

Five individuals, comprising two adults, an adolescent and two children, entered the cave barefoot and with a set of Pinus t. sylvestris/mugo bundles which were burned for illumination purposes. The adopted lighting system of several sticks enabled a longer period of lighting, as inferred from the fire wood illumination bundles adopted by the Bronze age salt miners at Hallstatt (Grabner et al., 2007; Grabner et al., 2010). Lighting bundles were usually made of resinous wood (Scots pine or Mountain pine), and called torch wood (Ast, 2001; Théry-Parisot et al., 2018). This interpretation fits with the archaeological evidence from the Bàsura Cave.

After a walk of approximately 150 m from the original opening of the cave and a climb of about 12 m, the group arrived at the ‘Corridoio delle impronte’. They proceeded roughly in single file, with the smallest individual behind, and walked very close to the side wall of the cave, a safer approach also used by other animals (e.g. Canidae incertae sedis and bears) when moving in a poorly lit and unknown environment. The slope of the tunnel floor, inclined by about 24°, may have further forced the individuals to proceed along the only flat area in the lower corridor, a couple of meters from the left wall of the cave. About 10 m from the ‘Corridoio delle impronte’, the cave roof drops to below 80 cm and members of the group were forced to crawl (Figure 11B), placing their hands (Figure 12) and knees (Figure 8b) on the clay substrate (Figure 9) (see also Video 1).

Figure 11

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Figure 12

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Video 1

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After a few meters, the group leader stopped, impressing two parallel calcigrade footprints, possibly to decide on the next movement and proceeded to cross the parts where the cave roof was at its lowest. The other individuals also stopped at the same place as the leader, and then proceeded along the same path by crawling and following the group leader, as indicated by the timing reconstructed from interactions between the tracks (Figure 9, d3).

After passing a bottleneck of blocks and stalagmites, the party descended for about ten meters along a steeply sloping surface. The whole group traversed a small pond, leaving deep tracks on the plastic waterlogged substrate, climbed a slope of 10 m beyond the ‘Cimitero degli orsi’, and finally arrived at the terminal room ‘Sala dei Misteri’, where they stopped. On the walls, several charcoal traces, generated by the torches, are preserved.

Some charcoal handprints produced by a flexed reaching up more than 170 cm on the roof of the ‘Sala dei Misteri’ confirm that the tallest individuals (Morphs. 4 and 5) were able to touch this part of the gallery. The fact that their footprints are not preserved relates to the loss of the central portion of the hall floor. In the same room, the adolescent and children started collecting clay from the floor and smeared it on a stalagmite at different levels according to height, as suggested by the breadth and relative distribution of the finger flutings on the karst structure. During their sojourn in the innermost room of the cave the young individual, that produced Morph. 2, imprinted ten clear heel traces (Citton et al., 2017), which are here interpreted as calcigrade tracks produced by a trackmaker who is momentarily standing-still to excavate and manipulate clay as was also recorded for the ‘Salle des Talons’ at Tuc d’Audoubert cave (Pastoors et al., 2015).

After stopping for several minutes (considering the quantity and ubiquity of the tracks), they exited and followed a route which did not always adhere to that followed on entry. After passing the small pond, they crossed the upper corridor following a more comfortable and safer route (Figure 11C). It is important to note that in the upper corridor all the prints point in the direction of the exit (Figures 11C and 13) while in the lower corridor, with the axis of the foot oriented parallel to the walls, most of the footprints are directed toward the interior of the cave.

Figure 13

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All the data are included in tables within the main text.

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Thank you for submitting your article "A multidisciplinary approach to a unique Palaeolithic human ichnological record from Italy (Bàsura Cave)" for consideration by eLife. Your article has been reviewed by three peer reviewers, including Jessica C. Thompson as the Reviewing Editor and Reviewer #1, and the evaluation has been overseen by Ian Baldwin as the Senior Editor. The following individuals involved in review of your submission have agreed to reveal their identity: Ignacio Díaz Martínez (Reviewer #2); Julien Riel-Salvatore (Reviewer #3).

The reviewers have discussed the reviews with one another and the Reviewing Editor has drafted this decision to help you prepare a revised submission.

Summary:

This manuscript reports a singular ichnological record left by a group of people who explored the Bàsura Cave around 14 thousand years ago. The authors complete a multidisciplinary study that complements the ichnological one based mainly on sedimentology, geochemistry, archaeobotany, geometric morphometrics, and 3D-modeling. They described footprints, handprints, knee and digit traces, etc. along two corridors and a cave chamber. The authors were able to identify 5 morphotypes of human footprints, which related to five ontogenetic states, two adults, an adolescent, and two children. The data and the methodology that the authors used in this study were high-quality and comprehensive: laser scanner and photogrammetrical 3D models of the cave and the trampling surfaces; new geochronological data; multivariate statistical analysis; biomorphic and biomechanical equations, etc. Therefore, the reconstruction proposed by the authors is based on solid arguments.

This is an excellent, thorough study on important material from an important site. It further uses innovative and cutting-edge methods (notably morphometrics) to reach new, original conclusions about human behavior in the Upper Paleolithic, shedding new light on what is essentially a snapshot of life at that time. The paper makes a real contribution to our understanding of life ca. 14,000 years ago. The combined findings about the number of individuals who visited the cave, their likely ages and the mode of their locomotion during their exploration are major contributions, as is the description of some of the behaviors in Sala dei Misteri. To this we can add the inference about the lack of footwear or clothing over the part of the bodies that left traces in the cave, which is instructive in its own right. The images are appropriate, convincing, and well-executed, and the video offers considerable vividness to their reconstruction. This work should finally give Grotta della Bàsurathe exposure in the international paleoanthropological community which it richly deserves by virtue of its uniqueness and remarkable degree of preservation.

Essential revisions:

1) Details of the dating: What is the evidence that this set of tracks represented a single event, rather than multiple events by different people over several visits? The close 14C dates on charcoal are compelling, but there is little context provided as to where the charcoal was recovered relative to the tracks. In the methods, the exact place from which the samples were derived relative to the marks should be made clear. There are data such as the square number etc. provided for each sample, but without knowing how the excavations and samples were positioned relative to the traces, it is unclear how they relate. Also, where the 14C dates are reported in the body of the manuscript, they should be indicated as being uncalibrated 14C dates BP (right now they don't specify). This becomes confusing where in the text, the authors suggest that the human explored the cave about 14,000 years ago, but the new data of the charcoal remains (Table 1) are around 12,000 years ago.

2) Details and discussion of the canid tracks: What is the significance of the canid tracks? With ongoing interest in the domestication of the dog, and the strong evidence that this process began in the Paleolithic, what are the implications of these tracks being interspersed with human tracks? Are they continuously associated with the human tracks, or do they appear incidental, as do the cave bear tracks? Figure 3 shows some tantalizing patterns, but they are not discussed further. This would be unusual behavior for most canids to enter the dark zone of a deep cave, although denning cannot be ruled out. Were there also canid bones as well as ursids? In the absence of such evidence, the authors should make an effort to establish why they do or do not believe the canid tracks are actually associated with the human entry. This could potentially be one of the more significant findings of the paper, but it is barely mentioned.

3) Details of the methods for trace recording, and which traces still remain in the cave: The methods are not quite detailed enough to establish the basis for the original data collection. Many of the figures describe casts from 1950, and this is noted in the methods, but then there are also parts of the methods that refer to tracing prints in situ. Are all of the traces now gone, or just some of them? How reliable are the casts likely to be? How much of the original traces are likely preserved in the casts? Are the positions of the traces as noted in the cave based only on casts, or on a combination of casts, historical notes, and modern remaining traces? If possible, each print described in Table 4 should have a descriptor saying where in the cave it is from and if it was in situ or a cast.

4) Details of other findings/context from the cave: The manuscript says Bàsura Cave contains some of the most significant human Paleolithic discoveries in Italy, and citations are provided, but a summary of what those discoveries are would be useful. What are they? How old are they? How do they likely relate to the activities recorded in the deeper parts of the cave?

5) Discussion of the symbolic context of the traces: What is the evidence that the finger flutings and finger traces shown in Figure 2 were created in a symbolic context? The presence of humans at the rear of the cave might suggest such behavior, but some finger traces themselves may be accidental, especially if they co-occur with the knee marks and suggest crawling. Perhaps those inferred to be symbolic should be in a separate figure from those inferred to derive from locomotion? The flutings appear significant, but without a bit more description of them it is difficult to discern which ones are actually on the walls/stalactites and which ones are just incidental and/or located on the floor. Also, what is the evidence that clay was collected from the floor? Are there places where this excavation seems to have taken place? The citations direct the reader to some of the primary descriptions, but a summary would be invaluable here.

6) Restriction of discussion to the evidence: In the discussion, there are some areas when the manuscript over-interprets the data. How do the trackways cause the inference that body sizes were slender and muscular? Is this based mostly on body mass estimates? The knee impressions do not seem to speak to this, since they are a part of the anatomy that has protruding bones and ligaments when kneeling in spite of the degree of slenderness and muscularity. Also, the ensuing interpretation that this was a hyper-active teenage explorer is an over-interpretation of the data and should be removed. It distracts from the more solid findings.

7) Revision of the discussion about the role of children: Another important conclusion of this study is that even very young children were included in potentially dangerous activities at the time. In that light, however, the authors should nuance their discussion of local comparative data from the Arene Candide and Arma Veirana, because all these show is that children were considered important members of their groups rather than saying anything about how they participated in their societies. In this sense, perhaps a better indirect referent for the social place of children in Epigravettian groups would be the small size of the pebbles argued to have been broken as part of mortuary behavior at the Arene Candide argued by Gravel-Miguel. I think this speaks more about the involvement of children as opposed to simply mentioning they were interred, which shows they were there but says little about their role.

8) Ichnological terminology: The authors used footprint, traces, tracks, etc. However, for all the ichnological record sensu lato, trace is the best option. If the trace is impressed by the foot, footprint, if it is impressed by the hand, handprint, etc.

9) Copy-editing: The paper needs to be thoroughly edited for the quality of its English. There are currently many odd turns of phrase and instances of transliteration from Italian to English that can be a bit jarring and impede smooth reading.

Essential revisions:

1) Details of the dating: What is the evidence that this set of tracks represented a single event, rather than multiple events by different people over several visits? The close 14C dates on charcoal are compelling, but there is little context provided as to where the charcoal was recovered relative to the tracks. In the methods, the exact place from which the samples were derived relative to the marks should be made clear. There are data such as the square number etc. provided for each sample, but without knowing how the excavations and samples were positioned relative to the traces, it is unclear how they relate. Also, where the 14C dates are reported in the body of the manuscript, they should be indicated as being uncalibrated 14C dates BP (right now they don't specify). This becomes confusing where in the text, the authors suggest that the human explored the cave about 14,000 years ago, but the new data of the charcoal remains (Table 1) are around 12,000 years ago.

In Figure 1 we have added the original position of the charcoal sample used for dating. The charcoal remains are those of the bundles originally utilized to illuminate the cave and preserved on the trampling palaeosurface of the “Sala dei Misteri”.

We add a new paragraph in the text highlighting all the evidences that make a single exploratory event for the cave more parsimonious, and the only one possible on the base of the available data. In particular, the short radiometric interval documented by both previous radiometric data and those derived from the latest research here presented (Table 1), together with the superimpositions between footprints allowing to decipher the timing of each singular impression, suggest that all individuals entered the cave in a single episode. In particular, the group of footprints C34, C35b, C36, C35 in the terminal part of the corridor of the footprints provides the timing of the impression: C35 (morph4) is superimposed on C36 (morph3) and the latter by C35 (morph4) (Figure 8). In the initial part of the same corridor it is observed that C63 (morph4) is superimposed by C64 (morph3) proving that morphs 3 and 4 are certainly contemporary. It is also difficult to assume that morphs 1 and 2 may have entered the deepest part of the cave without the presence of some older individual: if the older individuals who were leading the exploration were either morphs 3 or 4, the group consequently increases to include at least 4 individuals. No relationship can be established for the larger individual; however, as reported below, the fact that the footprints of this individual are regularly trampled and followed exactly on its path by all the other individuals, suggests that the larger one performed very likely the function of leader during a synchronous exploration. Therefore, the set of available evidences, and in particular the complex interference between the studied footprints, strongly suggests that a single exploration event by a heterogeneous restricted group of five individuals represents the most parsimonious and better supported hypothesis.

2) Details and discussion of the canid tracks: What is the significance of the canid tracks? With ongoing interest in the domestication of the dog, and the strong evidence that this process began in the Paleolithic, what are the implications of these tracks being interspersed with human tracks? Are they continuously associated with the human tracks, or do they appear incidental, as do the cave bear tracks? Figure 3 shows some tantalizing patterns, but they are not discussed further. This would be unusual behavior for most canids to enter the dark zone of a deep cave, although denning cannot be ruled out. Were there also canid bones as well as ursids? In the absence of such evidence, the authors should make an effort to establish why they do or do not believe the canid tracks are actually associated with the human entry. This could potentially be one of the more significant findings of the paper, but it is barely mentioned.

In fact, considering the importance of dog’s footprints as evidence for the domestication of the dog in the Paleolithic, this material is currently still under study and the object of a separate dedicated project; the latter includes the analysis of numerous trackways of extant dogs, hyenas and wolves to test whether the Bàsura dog footprints can be morphometrically referred to a single individual, and a detailed study of the interferences with the human footprints, to verify the real contemporaneity of frequentation of the hypogeal environment by dogs and humans. In any case, we included a short sentence in the new manuscript, along with some related bibliography, highlighting the possible importance of this material in the framework of dog domestication in the Paleolithic, stressing that this material is the subject of still ongoing study.

3) Details of the methods for trace recording, and which traces still remain in the cave: The methods are not quite detailed enough to establish the basis for the original data collection. Many of the figures describe casts from 1950, and this is noted in the methods, but then there are also parts of the methods that refer to tracing prints in situ. Are all of the traces now gone, or just some of them? How reliable are the casts likely to be? How much of the original traces are likely preserved in the casts? Are the positions of the traces as noted in the cave based only on casts, or on a combination of casts, historical notes, and modern remaining traces? If possible, each print described in Table 4 should have a descriptor saying where in the cave it is from and if it was in situ or a cast.

We add a paragraph in the Material and methods section stressing that in our direct analysis in the site we have also considered and analyzed the original casts from the 50s. Their position has been identified in the cave and it has been verified that all the footprints identified during the first explorations are still visible in situ (Figure 1). Some of them have suffered alterations and loss of details, others have been partially damaged. For this reason, in some cases we have integrated the morphometric data of the original in situ with those of plaster casts.

4) Details of other findings/context from the cave: The manuscript says Bàsura Cave contains some of the most significant human Paleolithic discoveries in Italy, and citations are provided, but a summary of what those discoveries are would be useful. What are they? How old are they? How do they likely relate to the activities recorded in the deeper parts of the cave?

In this regard we add some additional paragraphs in the Introduction, highlighting chronologically the first studies and published contributions to the Bàsura Cave immediately after the discovery. The historical overview, reported with sufficient detail, but without overloading to much the new version of the text, highlights the major contributions made by Tongiorgi, Lambroglia, Chiappella, Blanc and Pale, with the original interpretation of the human footprints, number of individuals and ritual interpretation of some traces and evidences.

5) Discussion of the symbolic context of the traces: What is the evidence that the finger flutings and finger traces shown in Figure 2 were created in a symbolic context? The presence of humans at the rear of the cave might suggest such behavior, but some finger traces themselves may be accidental, especially if they co-occur with the knee marks and suggest crawling. Perhaps those inferred to be symbolic should be in a separate figure from those inferred to derive from locomotion? The flutings appear significant, but without a bit more description of them it is difficult to discern which ones are actually on the walls/stalactites and which ones are just incidental and/or located on the floor. Also, what is the evidence that clay was collected from the floor? Are there places where this excavation seems to have taken place? The citations direct the reader to some of the primary descriptions, but a summary would be invaluable here.

Also, in this case, we are conducting a work totally dedicated to the analysis of the authors and age of the finger flutings, with 3D reconstruction of the “zoomorphic stalagmite” and identification of the possible source area from which the clay material was excavated. A figure has been added where a sample of involuntary types from those created in a possible symbolic context is differentiated.

6) Restriction of discussion to the evidence: In the discussion, there are some areas when the manuscript over-interprets the data. How do the trackways cause the inference that body sizes were slender and muscular? Is this based mostly on body mass estimates? The knee impressions do not seem to speak to this, since they are a part of the anatomy that has protruding bones and ligaments when kneeling in spite of the degree of slenderness and muscularity. Also, the ensuing interpretation that this was a hyper-active teenage explorer is an over-interpretation of the data and should be removed. It distracts from the more solid findings.

Yes, the slender and muscular body have been inferred on the base of body size estimate starting from footprint parameters. In this regard we add the following sentence to the new version of the text: “Body mass estimates derived from footprints parameters suggest slender and muscular body size for all the trackmakers. Arch angle and footprint morphology suggests a possible male as trackmaker of the largest footprint group. For the morphotypes 1, 2, 3, 4 any inference of gender result possible although the presence of almost a female (morph.4?) it seems probable.”

As properly suggested, the mention to ‘hyper-active teenage explorer’ has been completely removed from the text.

7) Revision of the discussion about the role of children: Another important conclusion of this study is that even very young children were included in potentially dangerous activities at the time. In that light, however, the authors should nuance their discussion of local comparative data from the Arene Candide and Arma Veirana, because all these show is that children were considered important members of their groups rather than saying anything about how they participated in their societies. In this sense, perhaps a better indirect referent for the social place of children in Epigravettian groups would be the small size of the pebbles argued to have been broken as part of mortuary behavior at the Arene Candide argued by Gravel-Miguel. I think this speaks more about the involvement of children as opposed to simply mentioning they were interred, which shows they were there but says little about their role.

In this respect, we have underlined the fact that not only some young individuals were buried, but that they were instead the object of particular attention. Indeed, we have added a mention to the grave goods found in association with the newborn from Arma Veirana and references are made to the sepulchral rites brought to light at the Arene Candide cemetery, along with the related bibliography. In our opinion, the constant association of adults and young individuals both in the burials and in external activities underlines the complex organization of Epigravettian and Mesolithic social groups, where even very young individuals, such as a newborn, could be fully included in the band structure both sharing their own responsibilities with adults and making themself object of elaborate funeral rites.

8) Ichnological terminology: The authors used footprint, traces, tracks, etc. However, for all the ichnological record sensu lato, trace is best option. If the trace is impressed by the foot, footprint, if it is impressed by the hand, handprint, etc.

We prefer to use the general term ‘trace’ sensu lato, since the material is not represented only by footprints and handprint (e.g. traces of knee and proximal shin).

9) Copy-editing: The paper needs to be thoroughly edited for the quality of its English. There are currently many odd turns of phrase and instances of transliteration from Italian to English that can be a bit jarring and impede smooth reading.

The new version of the manuscript was kindly reviewed carefully by the native speaker Professor Bruce Rubidge, consistently improving the quality of the English.

Author details

1. Marco Romano

   Evolutionary Studies Institute (ESI), School of Geosciences, University of the Witwatersrand, Johannesburg, South Africa

   Contribution

   Conceptualization, Data curation, Software, Formal analysis, Investigation, Visualization, Methodology

   For correspondence

   marco.romano@uniroma1.it

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0001-7629-3872

2. Paolo Citton

   CONICET-Consejo Nacional de Investigaciones Científicas y Técnicas, Buenos Aires, Argentina

   Contribution

   Conceptualization, Investigation, Methodology

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-6503-5541

3. Isabella Salvador

   MUSE, Museo delle Scienze, Trento, Italy

   Contribution

   Conceptualization, Resources, Data curation, Software, Formal analysis, Validation, Investigation, Visualization, Methodology

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-1058-3994

4. Daniele Arobba

   Museo Archeologico del Finale, Finale Ligure Borgo, Italy

   Contribution

   Formal analysis, Investigation

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-6946-7579

5. Ivano Rellini

   Dipartimento di Scienze della Terra, dell’Ambiente e della Vita (DISTAV), Università degli Studi di Genova, Genoa, Italy

   Contribution

   Formal analysis, Investigation

   Competing interests

   No competing interests declared

6. Marco Firpo

   Dipartimento di Scienze della Terra, dell’Ambiente e della Vita (DISTAV), Università degli Studi di Genova, Genoa, Italy

   Contribution

   Formal analysis, Investigation

   Competing interests

   No competing interests declared

7. Fabio Negrino

   Dipartimento di Antichità, Filosofia, Storia (DAFIST), Università degli Studi di Genova, Genoa, Italy

   Contribution

   Validation, Methodology

   Competing interests

   No competing interests declared

8. Marta Zunino

   Grotte di Toirano, Toirano, Italy

   Contribution

   Funding acquisition, Investigation, Project administration

   Competing interests

   No competing interests declared

9. Elisabetta Starnini

   1. Soprintendenza Archeologia Belle Arti e Paesaggio per la Città Metropolitana di Genova e le province di Imperia, La Spezia e Savona, Genoa, Italy
   2. Dipartimento di Civiltà e Forme del Sapere, Università di Pisa, Pise, Italy

   Contribution

   Supervision, Funding acquisition, Project administration

   Competing interests

   No competing interests declared

10. Marco Avanzini

    MUSE, Museo delle Scienze, Trento, Italy

    Contribution

    Conceptualization, Data curation, Software, Formal analysis, Supervision, Investigation, Visualization, Methodology

    Competing interests

    No competing interests declared

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