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.

1. 1. Ashton N
   2. Lewis SG
   3. De Groote I
   4. Duffy SM
   5. Bates M
   6. Bates R
   7. Hoare P
   8. Lewis M
   9. Parfitt SA
   10. Peglar S
   11. Williams C
   12. Stringer C

   (2014) Hominin footprints from early pleistocene deposits at Happisburgh, UK

   PLOS ONE 9:e88329.

   https://doi.org/10.1371/journal.pone.0088329

   • PubMed
   • Google Scholar
2. 1. Ast H

   (2001)

   Kienspan Und Unschlitt, beleuchtungimländlichenalltag

   Kulturbeilage Zum Amtsblatt Der Bezirkshauptmannschaft Wiener Neustadt.

   • Google Scholar
3. 1. Bavdekar SB
   2. Sathe S
   3. Jani P

   (2006)

   Prediction of weight of indian children aged upto two years based on foot-length: implications for emergency areas

   Indian Pediatrics 43:125–130.

   • PubMed
   • Google Scholar
4. 1. Blanc AC

   (1960)

   Le palline d’argilla della Grotta della Bàsura

   Rivista Di Studi Liguri 26:9–25.

   • Google Scholar
5. Book

   1. Blanc AC
   2. Pales L
   3. Lamboglia N

   (1960)

   Le Vestigia Umane Nella Grotta Della Bàsura a Toirano

   Bordighera: Istituto Internazionale di Studi Liguri.

   • Google Scholar
6. 1. Cheng Y-N
   2. Holmes R
   3. Wu X-C
   4. Alfonso N

   (2009) Sexual dimorphism and life history of Keichousaurus hui (Reptilia: Sauropterygia)

   Journal of Vertebrate Paleontology 29:401–408.

   https://doi.org/10.1671/039.029.0230

   • Google Scholar
7. 1. Chiappella VG

   (1952)

   Orsi e uomini preistorici nella grotta della strega

   Rivista Del Comune 29:22–29.

   • Google Scholar
8. 1. Chinnery B

   (2004) Morphometric analysis of evolutionary trends in the ceratopsian postcranial skeleton

   Journal of Vertebrate Paleontology 24:591–609.

   https://doi.org/10.1671/0272-4634(2004)024[0591:MAOETI]2.0.CO;2

   • Google Scholar
9. 1. Citton P
   2. Romano M
   3. Salvador I
   4. Avanzini M

   (2017) Reviewing the upper pleistocene human footprints from the ‘Sala dei Misteri’ in the Grotta della Bàsura (Toirano, northern Italy) cave: An integrated morphometric and morpho-classificatory approach

   Quaternary Science Reviews 169:50–64.

   https://doi.org/10.1016/j.quascirev.2017.05.016

   • Google Scholar
10. 1. Clarke HH

    (1933)

    An objective method of measuring the height of the longitudinal arch in the foot examinations

    Research Quarterly. American Physical Education Association 4:99–107.

    • Google Scholar
11. 1. Corrain C

    (1977)

    I resti scheletrici della sepoltura epigravettiana del “Riparo Tagliente’’ in Valpantena (Verona)

    Bollettino Del Museo Civico Di Storia Naturale Di Verona 4:35–79.

    • Google Scholar
12. 1. De Lumley MA
    2. Giacobini G

    (1985)

    Le impronte di piedi umani

    Rivista Di Studi Liguri 51:362–366.

    • Google Scholar
13. 1. de Lumley H
    2. Vicino G

    (1984) New data concerning the dating and interpretation of human footprints present in the “Grotta della Basura” at Toirano (Savona, Northern Italy). Results of an international round table

    Journal of Human Evolution 13:537–540.

    https://doi.org/10.1016/S0047-2484(84)80007-X

    • Google Scholar
14. 1. Dingwall HL
    2. Hatala KG
    3. Wunderlich RE
    4. Richmond BG

    (2013) Hominin stature, body mass, and walking speed estimates based on 1.5 million-year-old fossil footprints at Ileret, Kenya

    Journal of Human Evolution 64:556–568.

    https://doi.org/10.1016/j.jhevol.2013.02.004

    • PubMed
    • Google Scholar
15. 1. Formicola V
    2. Frayer DW
    3. Heller JA

    (1990) Bilateral absence of the lesser trochanter in a late epigravettian skeleton from arene candide (Italy)

    American Journal of Physical Anthropology 83:425–437.

    https://doi.org/10.1002/ajpa.1330830404

    • PubMed
    • Google Scholar
16. 1. Forriol F
    2. Pascual J

    (1990) Footprint analysis between three and seventeen years of age

    Foot & Ankle 11:101–104.

    https://doi.org/10.1177/107110079001100208

    • PubMed
    • Google Scholar
17. 1. Fryar CD
    2. Gu Q
    3. Ogden CL

    (2012)

    Anthropometric reference data for children and adults: United States, 2007-2010

    Vital and Health Statistics. Series 11, Data From the National Health Survey 11:1–48.

    • Google Scholar
18. Book

    1. Giacobini G

    (2008)

    La Grotta della Bàsura e il "mito neandertaliano"

    In: Arobba D, Maggi R, Vicino G, editors. Toirano E La Grotta Della Bàsura. Conoscere, Conservare E Gestire Il Patrimonio Archeologico E Paleontologico. Bordighera: Ist. Studi Liguri. pp. 21–27.

    • Google Scholar
19. Book

    1. Giannotti S

    (2008)

    La grotta della Bàsura (Toirano): rilettura e aggiornamento dei dati archeologici

    In: Arobba D, Maggi R, Vicino G, editors. Toirano E La Grotta Della Bàsura. Conoscere, Conservare E Gestire Il Patrimonio Archeologico E Paleontologico. Bordighera: Ist. Studi Liguri. pp. 233–246.

    • Google Scholar
20. 1. Grabner M
    2. Klein A
    3. Geihofer D
    4. Reschreiter H
    5. Barth FE
    6. Sormaz T
    7. Wimmer R

    (2007) Bronze age dating of timber from the salt-mine at Hallstatt, Austria

    Dendrochronologia 24:61–68.

    https://doi.org/10.1016/j.dendro.2006.10.008

    • Google Scholar
21. Book

    1. Grabner M
    2. Klein A
    3. Reschreiter H
    4. Barth F-E

    (2010)

    In Mining in European History and Its Impact on Environment and Human Societies – Proceedings for the 1st Mining in European History-Conference of the SFB-HIMAT

    Anreiter P, editors. Innsbruck: University Press.

    • Google Scholar
22. 1. Grivas TB
    2. Mihas C
    3. Arapaki A
    4. Vasiliadis E

    (2008) Correlation of foot length with height and weight in school age children

    Journal of Forensic and Legal Medicine 15:89–95.

    https://doi.org/10.1016/j.jflm.2007.05.007

    • PubMed
    • Google Scholar
23. 1. Hammer Ø
    2. Harper DAT
    3. Ryan PD

    (2001)

    PAST: paleontological statistics software package for education and data analysis

    Palaeontologia Electronica 41:1–9.

    • Google Scholar
24. Book

    1. Hammer Ø

    (2013)

    PAST Paleontological Statistics Version 3.0: Reference Manual

    Oslo: University of Oslo.

    • Google Scholar
25. 1. Janssens L
    2. Giemsch L
    3. Schmitz R
    4. Street M
    5. Van Dongen S
    6. Crombé P

    (2018) A new look at an old dog: bonn-oberkassel reconsidered

    Journal of Archaeological Science 92:126–138.

    https://doi.org/10.1016/j.jas.2018.01.004

    • Google Scholar
26. 1. Kanchan T
    2. Menezes RG
    3. Moudgil R
    4. Kaur R
    5. Kotian MS
    6. Garg RK

    (2008) Stature estimation from foot dimensions

    Forensic Science International 179:241.e1–242.e5.

    https://doi.org/10.1016/j.forsciint.2008.04.029

    • Google Scholar
27. 1. Krishan K
    2. Sharma A

    (2007) Estimation of stature from dimensions of hands and feet in a north indian population

    Journal of Forensic and Legal Medicine 14:327–332.

    https://doi.org/10.1016/j.jcfm.2006.10.008

    • Google Scholar
28. 1. Lamboglia N

    (1960)

    Le vestigia umane Nella Grotta della bàsura a toirano

    Rivista Di Studi Liguri 26:1–5.

    • Google Scholar
29. 1. Ledoux L
    2. Fourment N
    3. Maksud F
    4. Delluc M
    5. Costamagno S
    6. Goutas N
    7. Klaric L
    8. Laroulandie V
    9. Salomon H
    10. Jaubert J

    (2017) Traces of human and animal activity (TrAcs) in Cussac cave (Le Buisson-de-Cadouin, Dordogne, France): Preliminary results and perspectives

    Quaternary International 430:141–154.

    https://doi.org/10.1016/j.quaint.2016.06.002

    • Google Scholar
30. 1. Lemtur M
    2. Rajlakshmi C
    3. Damayanti D

    (2017) Estimation of stature from percutaneous length of ulna and tibia in medical students of Nagaland

    JDMS 16:46–52.

    https://doi.org/10.9790/0853-1601074652

    • Google Scholar
31. 1. Lupo KD

    (2017) When and where do dogs improve hunting productivity? the empirical record and some implications for early upper paleolithic prey acquisition

    Journal of Anthropological Archaeology 47:139–151.

    https://doi.org/10.1016/j.jaa.2017.05.003

    • Google Scholar
32. 1. Maineri D

    (1985)

    La scoperta della grotta della bàsura a toirano. Atti della tavola rotonda “La Grotta preistorica della Bàsura”, Toirano, 11-13 novembre 1983

    Rivista Di Studi Liguri 51:315–320.

    • Google Scholar
33. 1. Malina RM
    2. Hamill PVV
    3. Johston FE
    4. Lemeshow S

    (1973)

    Selected body measurements of children 6-11 years: United States

    Vital Health Stat pp. 1–4.

    • Google Scholar
34. 1. Mallegni F
    2. Bertoldi F
    3. Manolis S

    (2000)

    Palaeobiology of two gravettian skeletons from veneri cave (Parabita, Puglia, Italy)

    Homo : Internationale Zeitschrift FüR Die Vergleichende Forschung Am Menschen 51:235–257.

    • Google Scholar
35. 1. Mallegni F
    2. Fabbri PF

    (1995) The human skeletal remains from the upper palaeolithic burials found in Romito cave (Papasidero, Cosenza, Italy)

    Bulletins Et Mémoires De La Société d'anthropologie De Paris 7:99–137.

    https://doi.org/10.3406/bmsap.1995.2413

    • Google Scholar
36. 1. Molleson TI
    2. Oakley KP
    3. Vogel JC

    (1972) The antiquity of the human footprints of tana della basura

    Journal of Human Evolution 1:467–471.

    https://doi.org/10.1016/0047-2484(72)90078-4

    • Google Scholar
37. 1. Molleson T

    (1985)

    The antiquity of human footprints of tana della bàsura. Atti della tavola rotonda "La grotta preistorica della Bàsura, Toirano 11-13 novembre 1983

    Rivista Di Studi Liguri 51:367–372.

    • Google Scholar
38. 1. Morey DF
    2. Jeger R

    (2015) Paleolithic dogs: why sustained domestication then?

    Journal of Archaeological Science: Reports 3:420–428.

    https://doi.org/10.1016/j.jasrep.2015.06.031

    • Google Scholar
39. 1. Müller S
    2. Carlsohn A
    3. Müller J
    4. Baur H
    5. Mayer F

    (2012) Static and dynamic foot characteristics in children aged 1-13 years: a cross-sectional study

    Gait & Posture 35:389–394.

    https://doi.org/10.1016/j.gaitpost.2011.10.357

    • PubMed
    • Google Scholar
40. 1. Negrino F
    2. Benazzi S
    3. Hodgkins J
    4. Miller C
    5. Orr C
    6. Peresani M
    7. Riel-Salvatore J
    8. Strait D
    9. Gravel-Miguel C
    10. De Santis H
    11. Leger E
    12. Martini S
    13. Perroni E
    14. Laliberté A
    15. Pothier Bouchard G
    16. Starnini E
    17. Zerboni A

    (2017)

    On-going research and first data from middle and upper palaeolithic sites of Liguria region

    Bulletin Du Musée d'Anthropologie Préhistorique De Monaco 56:141–144.

    • Google Scholar
41. 1. Oberoi DV
    2. Kuruvilla A
    3. Saralaya KM
    4. Rajeev A
    5. Ashok B
    6. Nagesh KR
    7. Rao NG

    (2006)

    Estimation of stature and sex from footprint length using regression formulae and standard footprint length formulae respectively

    Journal of Punjab Academy of Forensic Medicine & Toxicology 6:5–8.

    • Google Scholar
42. 1. Pales L

    (1960)

    Le vestigia umane nella grotta della Bàsura a Toirano

    Rivista di Studi Liguri 26:9–90.

    • Google Scholar
43. 1. Paoli G
    2. Parenti R
    3. Sergi S

    (1980)

    Gli scheletri mesolitici della caverna delle arene candide (Liguria)

    Memorie dell'Istituto Italiano Di Paleontologia Umana Roma 3:33–154.

    • Google Scholar
44. 1. Pastoors A
    2. Lenssen-Erz T
    3. Ciqae T
    4. Kxunta U
    5. Thao T
    6. Bégouën R
    7. Biesele M
    8. Clottes J

    (2015) Tracking in caves: experience based reading of pleistocene human footprints in french caves

    Cambridge Archaeological Journal 25:551–564.

    https://doi.org/10.1017/S0959774315000050

    • Google Scholar
45. 1. Pastoors A
    2. Lenssen-Erz T
    3. Breuckmann B
    4. Ciqae T
    5. Kxunta U
    6. Rieke-Zapp D
    7. Thao T

    (2017) Experience based reading of pleistocene human footprints in Pech-Merle

    Quaternary International 430:155–162.

    https://doi.org/10.1016/j.quaint.2016.02.056

    • Google Scholar
46. 1. Pawar RM
    2. Pawar MN

    (2012) "Foot length--A functional parameter for assessment of height"

    The Foot 22:31–34.

    https://doi.org/10.1016/j.foot.2011.10.002

    • PubMed
    • Google Scholar
47. 1. Perri A

    (2016) A wolf in dog's clothing: Initial dog domestication and Pleistocene wolf variation

    Journal of Archaeological Science 68:1–4.

    https://doi.org/10.1016/j.jas.2016.02.003

    • Google Scholar
48. Book

    1. Riel-Salvatore J
    2. Gravel-Miguel C
    3. Maggi R
    4. Martino G
    5. Rossi S
    6. Sparacello VS

    (2018)

    New Insight into the Paleolithic Chronology and Funerary Ritual of Cavena delle Arene Candide

    In: Borgia V, Cristiani E, editors. Palaeolithic Italy – Advanced Studies on Early Human Adaptations in the Apennine Peninsula. Leiden: Sidestone Press. pp. 335–355.

    • Google Scholar
49. Book

    1. Robbins LM

    (1985)

    Footprints. Collection, Analysis, and Interpretation

    Charles C, editors. Illinois: Thomas Publisher.

    • Google Scholar
50. 1. Romano M

    (2017) Long bone scaling of caseid synapsids: a combined morphometric and cladistic approach

    Lethaia 50:511–526.

    https://doi.org/10.1111/let.12207

    • Google Scholar
51. 1. Romano M
    2. Citton P

    (2015) Reliability of digit length impression as a character of tetrapod ichnotaxobase: considerations from the Carboniferous-Permian ichnogenus Ichniotherium

    Geological Journal 50:827–838.

    https://doi.org/10.1002/gj.2601

    • Google Scholar
52. 1. Romano M
    2. Citton P

    (2017) Crouching theropod at the seaside. Matching footprints with metatarsal impressions and theropod authopods: a morphometric approach

    Geological Magazine 154:946–962.

    https://doi.org/10.1017/S0016756816000546

    • Google Scholar
53. 1. Ruff C
    2. Niskanen M
    3. Junno JA
    4. Jamison P

    (2005) Body mass prediction from stature and bi-iliac breadth in two high latitude populations, with application to earlier higher latitude humans

    Journal of Human Evolution 48:381–392.

    https://doi.org/10.1016/j.jhevol.2004.11.009

    • PubMed
    • Google Scholar
54. Conference

    1. Seitz SM
    2. Curless B
    3. Diebel J
    4. Scharstein D
    5. Szeliski R

    (2006) A comparison and evaluation of multi-view stereo reconstruction algorithms CVPR 2006

    2006 IEEE Computer Society Conference on Computer Vision and Pattern Recognition (CVPR'06). pp. 519–526.

    https://doi.org/10.1109/cvpr.2006.19

    • Google Scholar
55. 1. Shackelford LL

    (2007) Regional variation in the postcranial robusticity of late upper paleolithic humans

    American Journal of Physical Anthropology 133:655–668.

    https://doi.org/10.1002/ajpa.20567

    • PubMed
    • Google Scholar
56. 1. Sparacello VS
    2. Rossi S
    3. Pettitt P
    4. Roberts C
    5. Riel-Salvatore J
    6. Formicola V

    (2018) New insights on final epigravettian funerary behavior at arene candide cave (Western Liguria, Italy)

    Journal of Anthropological Sciences = Rivista Di Antropologia : JASS 96:1–24.

    https://doi.org/10.4436/JASS.96003

    • PubMed
    • Google Scholar
57. 1. Théry-Parisot I
    2. Thiébault S
    3. Delannoy J-J
    4. Ferrier C
    5. Feruglio V
    6. Fritz C
    7. Gely B
    8. Guibert P
    9. Monney J
    10. Tosello G
    11. Clottes J
    12. Geneste J-M

    (2018) Illuminating the cave, drawing in black: wood charcoal analysis at Chauvet-Pont d'arc

    Antiquity 92:320–333.

    https://doi.org/10.15184/aqy.2017.222

    • Google Scholar
58. Book

    1. Tongiorgi E
    2. Lamboglia N

    (1954)

    La Grotta Di Toirano

    Bordighera: Istituto Internazionale di Studi Liguri.

    • Google Scholar
59. Book

    1. Topinard P

    (1878)

    Anthropology

    London: Chapman and Hall.

    • Google Scholar
60. 1. Trinkaus E

    (1997) Appendicular robusticity and the paleobiology of modern human emergence

    PNAS 94:13367–13373.

    https://doi.org/10.1073/pnas.94.24.13367

    • PubMed
    • Google Scholar
61. 1. Ullman S

    (1979) The interpretation of structure from motion

    Proceedings of the Royal Society of London. Series B, Biological Sciences 203:405–426.

    https://doi.org/10.1098/rspb.1979.0006

    • PubMed
    • Google Scholar
62. 1. Villotte S
    2. Samsel M
    3. Sparacello V

    (2017) The paleobiology of two adult skeletons from baousso da Torre (Bausu da ture) (Liguria, Italy): Implications for gravettian lifestyle

    Comptes Rendus Palevol 16:462–473.

    https://doi.org/10.1016/j.crpv.2016.09.004

    • Google Scholar
63. 1. Walia S
    2. Modi BS
    3. Puri N
    4. Professor, Department of Anatomy, MMIMSR, Mullana, Ambala, Haryana, India

    (2016) Sexual dimorphism from foot dimensions and foot prints in haryanvi jat population

    International Journal of Anatomy and Research 4:2142–2147.

    https://doi.org/10.16965/ijar.2016.165

    • Google Scholar
64. 1. Yokoyama Y
    2. Guanjun S
    3. Nguyen Van H

    (1985)

    Dating of stalagmitic carbonates and bones of the bàsura cave at Toirano (Liguria, Italy) by the U-Th and U-Pa methods using alpha-and gamma-ray spectrometries

    Rivista Di Studi Liguri 51:373–378.

    • Google Scholar

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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