Sex Period. AC16. AC12. AC5. AC4. AC3. AC2. AC1_IP. 25000. 20000. 15000. 10000. 5000. 0. 25000. 20000. 15000. 10000. 5000. 0. J Humerus Left 35%. J H.
New data on Late Upper Paleolithic upper limb cross‐sectional geometry from Arene Candide: implications for Tardiglacial hunting practices VITALE S. SPARACELLO1,2, COLIN N. SHAW3 and DAMIANO MARCHI4. 1Archaeology, Durham University, 2Anthropology, University of New Mexico, 3McDonald Institute for
Archaeological Research, University of Cambridge, 4Department of Biology, University of Pisa
Introduction European Late Upper Paleolithic people display levels of bilateral asymmetry in humeral mechanical strength (calculated via cross‐sectional geometry, CSG; Ruff et al., 2006) comparable to professional tennis players (Churchill et al., 1996, 2000; Shackelford, 2007; Fig. 1). This highly‐ characteristic trait has been associated with the use of throwing weapons to “kill at a distance” (Churchill and Rhodes, 2009) This activity is not only strenuous, but requires continuous training from a young age to achieve both speed and accuracy (Cattelain, 1997; Whittaker and Kamp, 2006). The plausibility of this hypothesis was supported by experimental evidence (Schmitt et al., 2003), and by the finding of high asymmetry in humeral retroversion and medial humeral epicondyle enthesopathies in the dominant limb of LUP males, traits that are both associated with throwing in modern day athletes (Rhodes and Churchill, 2009; Villotte et al., 2010). However, previous studies have pooled European samples to increase sample size, which nevertheless remained low (Fig. 1). In addition, this pooling may have masked regional behavioral variation associated with the fragmentation of populations during the Tardiglacial period (ca. 16‐10,000 BP). During a recent re‐assessment of the skeletal elements in secondary deposition from Arene Candide (north‐western Italy, Fig. 2), we were able to collect the first CSG data from the upper limb, and obtained the largest sample of Late Paleolithic coupled elements coming from the same site. In addition, we identified further skeletal elements belonging to an individual with probable congenital rickets (AC3; Formicola, 1995).
Figure 1 – Humeral bilateral asymmetry (HUMBA) in groups practicing different activities
Materials and Methods Data consist in upper limb (humerus, radius, and ulna) CSG mechanical rigidity (J) and bilateral asymmetry from six adult males (AC 2, 3, 4, 5, 10, and 12) and one adolescent (13 y.o., AC 16). Solid CSG was calculated from subperiosteal contours following Sparacello and Pearson (2010). Data collection was performed via silicone molds and structured‐light 3D scanning (DAVID SLS‐2).
Results
Figure 2 – Left, the Arene Candide Epigravettian «necropolis»; Above, the juvenile AC 16 Formicola and Toscani, 2015.
All adult Epigravettian individuals show high levels of mid‐distal and midshaft humeral bilateral asymmetry (40% to 130%, Fig. 3‐4). Ulnar and radial asymmetry is not always consistent with humeral lateralization (Table 1). The adolescent individual displays low humeral (12%) and ulnar (16.5%) asymmetry. Conversely, the adult with diagnosed congenital x‐linked rickets displays high humeral asymmetry (40%). BARGR2
20000 J Humerus Right 35%
AC5 AC12 AC2
15000
AC1_IP
AC3 AC4
AC10
10000
AC16
Sex F F M M
5000
0
Fig. 3
0
5000
10000 15000 J Humerus Left 35%
Period EUP LUP EUP LUP
20000
25000
DOLV15 DOLV3 PAG25 DOLV13 DOLV14 DOLV16 GDENF4 PAVLO1 SUNGHIR1 CAPBL1 Rom_1 Rom_4 Rom_5 SANTEO 1 STGERIV4 AC1_IP AC2 AC3 AC4 AC5 AC10 AC12 AC16 CHANC1 GOUGH1 LAUGB4 NEUES2 Rom_3 Rom_6 Rom_7 Rom_8 ROMAN1 VILLABR
SEX
PERIOD
F F F M M M M M M F F F F F F M M M M M M M M M M M M M M M M M M
EUP EUP EUP EUP EUP EUP EUP EUP EUP LUP LUP LUP LUP LUP LUP EUP LUP LUP LUP LUP LUP LUP LUP LUP LUP LUP LUP LUP LUP LUP LUP LUP LUP
30000
HUMBA 35%
SEX PERIOD
34.96
25000
AC5
55.15
32.39
19.32 110.92 40.57 63.06 54.58 41.01 97.41 12.07
J Humerus Right 50%
25000
AC2
20000
AC12 AC4 AC3
AC1_IP
15000
10000 AC16
Sex F F M M
5000
56.25
0
Fig. 4
0
5000
10000 15000 20000 J Humerus Left 50%
Period EUP LUP EUP LUP
25000
Figure 3, 4 – Scatterplot of right vs left humeral bilateral asymmetry in mechanical rigidity (J). The line represent isometry.
30000
PAG25 PREDM10 GDENF4 PAVLO1 PREDM14 PREDM3 PREDM4 SUNGHIR1 AC1_IP AC2 AC3 AC4 AC5 AC12 AC16 CAPBL1 Rom_4 STGERIV4 CHANC1 GOUGH1 LAUGB4 ROMAN1
F F M M M M M M M M M M M M M F F F M M M M
EUP EUP EUP EUP EUP EUP EUP EUP EUP LUP LUP LUP LUP LUP LUP LUP LUP LUP LUP LUP LUP LUP
HUMBA 50% 13.54
25.31
49.09 128.31 86.45 72.00 73.93 92.65 13.87 29.21
51.59
Radius AC2 M AC4 M Ulna AC2 M AC3 M AC4 M AC12 M AC16 M
RADBA 20% 76.35 12.86 ULBA20% 27.69 4.26 42.50 53.99
RADBA50% 51.46 17.25 ULBA50% 57.87 4.28 5.91 48.21
RADBA66% 27.19 13.94 ULBA66% 37.73 7.06 15.23 31.56 16.37 Table 1 – Radial and ulnar bilateral asymmetry, calculated as BA=100*(Jmax‐ Jmin)/Jmin
Conclusions All adult males from Arene Candide show a level of humeral bilateral asymmetry that is comparable to the one seen in professional tennis players. This suggests that stressful unimanual activities (presumably throwing weapons during the hunt) were widely shared among all members of the group. This includes Arene Candide 3, whose presumably genetic disorder (possibly hypophosphatemic rickets; Formicola, 1995) resulted in low stature, diffused enthesopathies, and bowing of the diaphyses, especially in the right lower limb, which likely affected his ability to perform high‐mobility tasks (see the very curved fibula, and the asymmetry of L5 probably due to long‐term unbalanced gait; Fig 5). Yet, this individual is mechanically robust, and, like “his peers”, appears to have performed throwing. A similar result was found some 15,000 years before, and 800 km east, in the Gravettian Dolní Vĕstonice 15 (Trinkaus et al., 2001), which also shows bowing deformities presumably due to an hereditary condition. The high frequency of hereditary dysplasias, probably due to low genetic diversity caused by inbreeding, the special funerary treatment of the individuals showing signs of those conditions, and evidence of a society where persistence in the face of pathology was necessary, all suggest a strong parallel between the Gravettian and Epigravettian societies across time and space. Despite the expectations of an early onset of throwing behavior, and observations of high asymmetry in Gravettian juveniles (Sunghir; Cowgill et al., 2012), upper limb asymmetry in AC 16 is not particularly high. The absence of comparative individuals render difficult to determine whether this is due to chance, or to a later onset of throwing behavior when Figure 2 – Bowing deformities, enthesopathies, and asymmetry in the fifth lumbar of Arene Candide 3. compared to the Early Upper Paleolithic (see also the high humeral midshaft values of Research funded by the Marie Skłodowska‐Curie actions ‐ European Commission, and by the Wolfson Institute for Health and Wellbeing, Durham,UK asymmetry in Arene Candide 1, Il Principe). Cattelain P. 1997. Hunting during the Upper Paleolithic Bow Spearthrower or Both? In: Knecht H, editor. Projectile Technology. New York: Springer. p. 213‐240. Churchill SE, Rhodes J. 2009. The evolution of the human capacity for “killing at a distance”: the human fossil evidence for the evolution of projectile weaponry. In: Hublin J‐J, Richards MP, editors. The evolution of hominin diets. New York: Springer. p. 201‐210. Churchill SE, Weaver AH, Niewoehner WA. 1996. Late Pleistocene human technological and subsistence behavior: functional interpretations of upper limb morphology. Quaternaria Nova 6:413‐447. Churchill SE, Formicola V, Holliday TW, Holt BM, Schumann BA. 2000. The Upper Paleolithic population of Europe in an evolutionary perspective. In: Roebroeks W, Mussi M, Svoboda J, Fennema K, editors. Hunters of the Golden Age: the Mid Upper Paleolithic of Eurasia (30,000–20,000 bp). Leiden: Leiden University Press. p 31–57. Cowgill LW, Mednikova MB, Buzhilova AP, Trinkaus E. 2012. The Sunghir 3 Upper Paleolithic Juvenile: Pathology versus Persistence in the Paleolithic. Int J Osteoarch doi 10.1002/oa.2273. Formicola V. 1995. X‐linked hypophosphatemic rickets: a probable Upper Paleolithic case. Am J Phys Anthropol 98:403–409. Formicola V, Toscani F. 2015. A Late Epigravettian skeleton of an adolescent from Arene Candide cave. Bull Mus Anthropol Prehist Monaco. In press. Rhodes JA, Churchill SE. 2009. Throwing in the Middle and Upper Paleolithic: inferences from an analysis of humeral retroversion. J Hum Evol 56:1‐10.
Schmitt D, Churchill SE, Hylander WL. 2003. Experimental evidence concerning spear use in Neanderthals and Early Modern humans. J Archaeol Sci 30:103‐ 114. Shackelford LL. 2007. Regional variation in the postcranial robusticity of late Upper Paleolithic humans. Am J Phys Anthropol 133:655‐668. Sparacello VS, Pearson OM. 2010. The importance of accounting for the area of the medullary cavity in cross‐sectional geometry: a test based on the femoral midshaft. Am J Phys Anthropol 143:612‐624. Trinkaus E, Formicola V, Svoboda J, Hillson SW, Holliday TW. 2001. Dolní Vĕstonice 15: Pathology and persistence in the Pavlovian. J Archaeol Sci 28:1291‐ 1308. Villotte S, Churchill SE, Dutour OJ, Henry‐Gambier D. 2010. Subsistence activities and the sexual division of labor in the European Upper Paleolithic and Mesolithic: evidence from upper limb enthesopathies. J Hum Evol 59:35‐43. Whittaker JC, Kamp KA. 2006. Primitive weapons and modern sport: atlatl capabilities, learning, gender, and age. Plains Anthropol 198:213‐221.
