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Temporal changes in Arikara humeral and femoral cross- sectional geometry associated with horticultural intensification. Journal of Archaeological Science, vol.
ARCHAEOLOGY, ETHNOLOGY & ANTHROPOLOGY OF EURASIA Archaeology Ethnology & Anthropology of Eurasia 42/3 (2014) 141–156 E-mail: [email protected]

141

ANTHROPOLOGY S.S. Tur Altai State University, Leninsky Pr. 61, Barnaul 656049, Russia E-mail: [email protected]

BILATERAL ASYMMETRY OF LONG BONES IN BRONZE AND EARLY IRON AGE PASTORALISTS OF THE ALTAI*

The aim of the study is to explore patterns of directional asymmetry (DA) of long bones among the ancient pastoralists of the Russian Altai. Long bones of the upper and lower limbs and clavicles were measured bilaterally in two temporally diverse skeletal samples dating to the Middle Bronze Age and the Early Iron Age. Statistically signi¿cant sex and chronological differences were found in DA of the upper limb diaphyseal breadths, which are strongly inÀuenced by mechanical factors during life. These results suggest that manual loadings were bilaterally symmetric in males, but not in females. Sexual dimorphism in the upper-limb-use asymmetry was greater in the later group than in the earlier group. Besides, the female subgroups exhibited strong DA in features evidencing biomechanical stress on the femur. Temporal differences in DA of the upper limb length are possibly due to changes in the level of environmental and/or genetic stress. Keywords: Bilateral asymmetry, physical activity, long bones, pastoralists, Bronze Age, Early Iron Age, Scythian period, Altai.

Introduction The degree of bilateral asymmetry of human bones can be an indicator of biological adaptation to various factors of external and internal environment. There are three basic types of bilateral asymmetry: fluctuating (FA), directional (DA), and antisymmetry (AnS), which differ in the nature of differences between the right and left sides. FA indicates minor random differences that are normally distributed around zero mean. In DA, also described as *Supported by the Russian Foundation for Basic Research (Project No. 11-06-00360ɚ) and the Russian Foundation for the Humanities (Project No. 12-01-00159).

fixed asymmetry, one side is larger than the other on average (the differences are also normally distributed whereas the mean is signi¿cantly greater than zero). In AnS, or random asymmetry, the difference between sides is signi¿cant, but the larger side is determined randomly (the distribution of differences is bimodal or platykurtic with zero mean (Palmer, Strobeck, 1986, 2003)). Various types of asymmetry often interact. The most informative types in the study of ancient populations are FA and DA. FA measures the level of developmental instability, which increases under environmental and genetic stress (Harris, Nweeia, 1980; Livshits, Kobyliansky, 1991; Hershkovitz et al., 1993; Markow, Martin, 1993; Gray, Marlowe, 2002; Guatelli-Steinberg, Sciulli, Edgar, 2006;

Copyright © 2014, Siberian Branch of Russian Academy of Sciences, Institute of Archaeology and Ethnography of the Siberian Branch of the Russian Academy of Sciences. Published by Elsevier B.V. All rights reserved. doi:10.1016/j.aeae.2015.04.016

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Schaefer et al., 2006; DeLeon, 2007; Hoover, Matsumura, 2008; Graham et al., 2010; Özener, 2010). However, measuring FA under considerable DA is technically very difficult (Palmer, 1994; Graham et al., 1998), which imposes certain restrictions on the choice of traits. Studies of FA in ancient populations are normally based on dental and cranial characters. DA of human bones is related to functional asymmetry (motor, sensory, and psychic). Motor asymmetry may be related to unilateral motions, when mostly one (dominant) arm is used, to functional specialization of the legs, to the development of motor capacities, and to the conditional response speed. DA of long bones is largely caused by functional adaptation to mechanical factors. Relationship between DA and mechanical loading on limb bones, especially on those of the upper limbs, is widely used in the study of physical stress in ancient populations with regard to lifestyle and work, and of the sexual division of labor. Most such studies, however, focused on huntergatherers and farmers residing in regions other than Russia (Ruff, Jones, 1981; Bridges, 1989; Fresia, Ruff, Larsen, 1990; Stirland, 1993; Sakaue, 1998; Mays, 1999; Bridges, Blitz, Solano, 2000; Stock, Pfeiffer, 2004; Westcott, Cunningham, 2006; Sládek et al., 2007; Wanner et al., 2007; Kujanová et al., 2008; Maggiano et al., 2008; Sparacello, Marchi, 2008; Pomeroy, Zakrzewski, 2009; Weiss, 2009; Sparacello et al., 2011). The objective of the present study is to evaluate directional asymmetry of long limb bones and clavicles in Bronze and Early Iron Age pastoralists of the foreststeppe zone of the Altai. Materials and methods Long bones of upper and lower limbs and clavicles from burials associated with two cultures of the forest-steppe Altai were studied: Middle Bronze Age Andronovo Culture (AC) and Early Iron Age (Scythian era) Staroaleika Culture (SC). The economy of both groups was mainly pastoral. The AC sample includes skeletons from Barsuchikha-1, Zolotushka, Malopanyushevsky, Manzhikha-5, Teleutsky Vzvoz-1, Firsovo-14, Chekanovsky Log-2 and 10, whereas the SC sample consists of skeletons from Firsovo-14 and Tuzovskie Bugry-1. Collections are owned by the Altai State University Museum of Archaeology and Ethnography Department of Anthropology. Sex and age were estimated by standard methods. Age differences between the two samples are insigni¿cant. Long bones of arms and legs (humeri, ulnae, radii, femora, tibiae), and clavicles were measured on both sides. Bones with incompletely fused epiphyses as well as deformed and pathologically affected ones were excluded. Limb bones were oriented in the medio-lateral and sagittal planes according to the technique proposed by C. Ruff

(Ruff, Hayes, 1983; Ruff, 2002). The midshaft of tibiae was assessed on the basis of total length (Ɍ1) whereas that of other bones (H1, R1, U1, F1) was based on maximal length. The accuracy of longitudinal measurements was within 0.5 mm, and that of transverse diameters of the shaft, within 0.05 mm. Apart from standard measurements, numbered after R. Martin (Alekseyev, 1966), several additional ones were taken in the study of DA. These include the average midshaft diameter, calculated as a half-sum of the sagittal and medio-lateral diameters. Also midshaft diameters of humeri (sagittal, medio-lateral, and average) were taken at 35 % maximal length from the distal end. On the tibiae, minimal and maximal midshaft diameters were measured. To calculate the measurement error, long bones of seven individuals were measured three times with an interval of several weeks. The error was calculated by the method proposed by T.D. White (White, Folkens, 2005): deviations from the mean were averaged and divided by the mean. For most measurements, the error was less than 0.5 % and therefore could not have affected the results in an appreciable way (Table 1). For a qualitative evaluation of DA the standardized directional asymmetry coef¿cient, %DA, was calculated according to the formula: %DA =

,

R being the dimension on the right side, L, that on the left side This formula, which is widely used (Steele, Mays, 1995; ýuk, Leben-Seljak, Štefanþiþ, 2001; Mays, 2002; Auerbach, Ruff, 2006; Blackburn, Knüsel, 2006; Sládek et al., 2007; Auerbach, Raxter, 2008; Kujanová et al., 2008; Jaskulska, 2009; Pomeroy, Zakrzewski, 2009; Weiss, 2009; Stock et al., 2013), makes it possible to compare the asymmetry of dimensions regardless of their absolute value as in the case of bone length and shaft breadth. The %DA value is positive if the right dimension is larger, and negative in the opposite case. The normality of the %DA distribution was assessed with the Lilliefors test. If no significant departures from normality were found, the one-sample t-criterion was applied to test if the mean %DA significantly differs from zero. In case the distribution of %DA was signi¿cantly non-normal, the Wilcoxon nonparametric statistics was used to test the signi¿cance of difference between the sides. To evaluate the sexual and temporal differences, the nonparametric Mann-Whitney U-test was employed. For calculating the proportion of individuals with the predominance of the right and left side, only %DA > 0,5 %, were used to avoid the effect of measurement error and FA (Auerbach, Ruff, 2006). Signi¿cance of between-

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Table 1. Measurement errors (n = 7), % Traits*

Right

Left

Average

C1

0.11

0.13

0.12

C4

0.77

0.84

0.81

C5

1.20

0.89

H1

0.05

H4

Traits

Right

Left

Average

R5a

0.50

0.41

0.46

F1

0.04

0.05

0.05

1.05

F6

0.36

0.23

0.30

0.04

0.05

F7

0.05

0.15

0.10

0.28

0.33

0.31

F21

0.21

0.21

0.21

H6b

0.29

0.17

0.23

T1

0.05

0.09

0.07

H6c

0.25

0.35

0.30

T5

0.28

0.11

0.20

R1

0.05

0.06

0.06

T8

0.52

0.57

0.55

R4a

0.39

0.59

0.49

T9

0.51

0.85

0.68

*See Table 2 for explanations.

group differences in these coef¿cients was assessed by the chi-squared test. The correlation between the DA of longitudinal and transverse measurements was evaluated with the Pearson correlation coef¿cient using regression residuals of logtransformed dimensions on the right and left side (Ibid.). All calculations were conducted using the STATISTICA 10 software. To assess the between-group variation of %DA values for humeri, radii, femora, and tibiae, data relating to nine groups of vastly different origin, living in various parts of the world, were taken from the study by Auerbach and Ruff (2006). Also, data on %DA of humeri and clavicles of 15 more groups were taken from the study by Auerbach and Raxter (2008). Most of those groups are recent. Results Upper limbs and clavicles. No DA of clavicular length was found in either the male or the female AC samples. Arm bones show a slight right-side predominance, but only the %DA of humeri in the pooled sample is statistically signi¿cant (Table 2). In the SC samples (male, female, and pooled), the right humeri, radii, and ulnae are longer than their left counterparts; the right clavicles, by contrast, are shorter (table 3). Bronze Age and Scythian Age groups differ in the %DA of humeral length (males: ɪ = 0.035; females: ɪ = 0.011; pooled, ɪ = 0.009) and of ulnar length (females: ɪ = 0.007; pooled: ɪ = 0.028) (¿g. 1). In the SC sample, the proportion of individuals with the rightside predominance of humeral length is higher (pooled, ɪ = 0.021), whereas the frequency of individuals with the left-side predominance is lower (females, ɪ = 0.020; pooled, ɪ = 0.025) (Table 4). The DA of transverse dimensions is generally more variable than that of bone length. In SC females, the shaft

of clavicle shows a marked right-bias. In AC samples, the respective values of %DA demonstrate increased variation, females showing a left-bias. Among the ten transverse dimensions of the humeral shaft, seven in the AC sample and four in SC sample show a signi¿cant right-bias. In all groups, the sagittal midshaft diameter H6C demonstrates the highest positive %DA scores regardless of sex. Not only are absolute measurements signi¿cantly asymmetric, but so are coef¿cients of midshaft cross-section H6 : H5 and H6c : H6b (Table 5), and the same is true of the proximal and distal epiphyseal breadth. The asymmetry of longitudinal diameters of humeri is significantly correlated with that of the transverse diameters (Table 6). Radial shafts show signi¿cant right-bias in R4 and R4a breadth. In AC samples, the %DA of the midshaft sagittal diameter of R5a is signi¿cantly negative. The midshaft cross-section indices R5 : R4 and R5a : R4a, too, reveal considerable asymmetry in both male and female groups. The distal epiphyseal breadth of radii is less asymmetric than the shaft width. The highest %DA score in ulnar shafts are those of width, exhibiting left-bias except for the female SC sample. When the %DA scores of transverse dimensions of clavicles and arm bones in male and female Bronze Age and Early Iron Age series are compared, signi¿cant contrasts turn up. Sex differences concern the asymmetry of clavicular shafts, speci¿cally the vertical diameter C4 (ɪ = 0.048) in the AC sample, and C6 circumference (p = 0.043) in the SC sample. Humeral shafts are more asymmetric in females. In the AC group, sexes signi¿cantly differ in the %DA score of the midshaft sagittal diameter H6c (ɪ = 0.040), in the H6c : H6b ratio (ɪ = 0.049). In the SC group, sex differences concern the sagittal diameter at the 35 % length level (ɪ = 0.036) and midshaft circumference H7a (ɪ = 0.043). In addition, Early Iron Age females have a

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Table 2. Directional asymmetry indexes of clavicles and limb bones in the Andronovo sample Trait 1

n1/n2 2

%DA Males

Females

Pooled

3

4

5

Clavicula C1. Length

14/9

–0.62 (0.13)

–0.34 (–0.41)

–0.37 (0.08)

C4. Vertical diameter

15/9

2.62 (1.84)

–2.47 (–5.38)

1.46 (–0.87)

C5. Sagittal diameter

14/9

–0.60 (0.12)

–0.84 (1.26)

–0.80 (0.56)

Mean diameter (CAD)

14/9

–0.10 (0.91)

–2.60 (–1.94)

–0.43 (–0.21)

C6. Circumference

14/9

0.02 (0.69)

–1.55 (–1.51)

–1.29 (–0.17)

Humerus H1. Maximal length

19/21

0.58 (0.47)

0.81 (0.48)

0.65 (0.47) ɚ

H3. Upper epiphysis breadth

11/13

0.00 (0.55)

2.04 (1.43) b

0.99 (1.02) ɚ

H4. Lower epiphysis breadth

17/16

0.00 (0.80) ɚ

1.69 (1.89) c

1.48 (1.33) c

H5. Maximal diameter of midshaft

21/23

0.95 (1.22)

2.37 (2.24) b

2.02 (1.75) c

H6. Minimal diameter of midshaft

21/23

0.00 (–0.59)

–0.62 (–1.28)

–0.26 (–0.95)

H6b. Midshaft breadth

21/22

–0.42 (–0.48)

–1.90 (–1.55)

–1.09 (–1.03)

H6c. Sagittal diameter of midshaft

21/23

0.84 (2.08)

Average diameter of midshaft (HAD)

21/22

Shaft breadth at 35 % length (H35 % ml)

c

3.12 (3.57) c

0.21 (0.78)

1.54 (1.62) ɚ

0.93 (1.21) ɚ

18/19

0.95 (0.50)

1.19 (1.09)

1.19 (0.80)

Sagittal diameter at 35 % length (H35 % ap)

18/19

1.34 (1.05)

0.51 (0.59)

1.09 (0.81) b

Mean diameter at 35 % length (HAD35 %)

18/17

0.71 (0.78)

0.77 (1.01)

0.77 (0.89) ɚ

H7. Minimal circumference of shaft

17/15

0.72 (0.54)

0.87 (1.09)

0.82 (0.80) ɚ

H7a. Midshaft circumference

20/21

0.73 (0.71)

0.81 (1.02)

0.74 (0.87) ɚ

14/17

–0.69 (–0.27)

–0.47 (0.02)

–0.47 (–0.11)

0.38 (0.05)

0.42 (0.61)

0.38 (0.27)

3.78 (3.71) ɚ

5.57 (4.91) c

H10. Vertical diameter of head

ɚ

4.38 (4.94)

Radius R1. Maximal length

14/9

b

R4. Maximal breadth of midshaft

16/14

6.03 (5.97)

R5. Sagittal diameter of shaft at maximal breadth level

16/14

–0.58 (–0.55)

0.46 (1.44)

0.00 (0.38)

Mean diameter of shaft at the maximal breadth level (RAD max)

16/14

3.19 (3.22) ɚ

3.40 (2.74) b

3.19 (2.99) c

R4a. Midshaft breadth

15/15

7.82 (8.33) c

4.43 (3.39)

5.34 (5.86) c

R5a. Sagittal diameter of midshaft

15/15

–3.92 (–4.37) b

–1.90 (–1.16)

–2.87 (–2.76) b

Mean diameter of midshaft (RAD)

15/18

3.18 (2.81) b

1.41 (1.43)

2.64 (2.12) ɚ

R3. Minimal shaft circumference

14/15

1.07 (0.65)

1.26 (1.44) ɚ

1.24 (1.06) ɚ

16/16

0.51 (0.71)

0.78 (0.54)

0.72 (0.62)

0.64 (0.39)

0.00 (0.16)

0.28 (0.31)

Breadth of distal epiphysis (R dist)

Ulna U1. Maximal length

18/10

U11. Sagittal diameter of shaft at the maximal breadth level

19/13

–0.70 (–0.58)

–3.05 (0.21)

–2.48 (–0.26)

U12. Maximal shaft breadth

20/13

–1.32 (–1.15)

–4.42 (–2.13)

–1.95 (–1.54)

Mean diameter at the maximal breadth level (UAD max)

19/13

–1.51 (–0.83)

–2.21 (–1.04)

–1.60 (–0.92)

S.S. Tur / Archaeology, Ethnology and Anthropology of Eurasia 42/3 (2014) 141–156

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Table 2 (end) 1

2

3

4

5

U11a. Sagittal diameter of midshaft

15/11

–1.55 (–1.35)

–2.30 (1.11)

–1.92 (–0.31)

U12a. Midshaft breadth

15/11

–2.09 (–3.51)

–7.58 (–5.09)

–3.64 (–4.18) ɚ

Mean diameter of midshaft (UAD)

15/11

–4.17 (–2.49) ɚ

–4.20 (–2.18)

–4.18 (–2.36) ɚ

U3. Minimal shaft circumference

18/16

–0.59 (–0.28)

–0.70 (–0.58)

–0.59 (–0.42)

Femur F1. Maximal length

26/22

–0.11 (–0.12)

–0.48 (–0.36) ɚ

–0.22 (–0.23)

F6. Sagittal diameter of midshaft

28/22

0.17 (0.59)

–0.58 (–0.53)

0.00 (0.10)

F7a. Transverse diameter of midshaft

27/22

–1.66 (–1.24)

Mean diameter of midshaft (FAD)

27/22

F8. Midshaft circumference

b

–1.88 (–2.05) c

–0.71 (–0.17)

–1.73 (–1.78) b

–0.79 (–0.97) ɚ

28/21

–0.54 (–0.37)

–0.59 (–1.37)

–0.54 (–0.80) ɚ

F21. Lower epiphysis breadth

17/16

0.00 (0.01)

0.00 (0.13)

0.00 (0.07)

F18. Caput height

16/15

0.86 (0.74)

0.67 (0.74)

0.67 (0.74) ɚ

–0.28 (–0.48) ɚ

–0.28 (–0.27)

–0.28 (–0.38) b

ɚ

–2.74 (–3.03)

Tibia T1. Full length

23/19

T5. Maximal breadth of upper epiphysis

11/8

0.60 (0.46)

0.34 (0.16)

0.60 (0.34)

T6. Maximal breadth of lower epiphysis

14/9

–0.09 (–0.30)

0.00 (0.93)

0.00 (0.18)

T8. Sagittal diameter of midshaft

24/16

–0.34 (–0.38)

–1.29 (–1.96)

–0.94 (–1.01)

T8a. Sagittal diameter at the level of nutrient foramen

22/18

–2.60 (–1.61)

–3.04 (–2.81) ɚ

–2.91 (–2.15) b

T9. Transverse diameter of midshaft

24/16

1.13 (2.19) ɚ

3.94 (4.18) b

1.65 (2.98) b

T9a. Transverse diameter of shaft at the level of nutrient foramen

22/17

–0.41 (–0.07)

2.14 (2.88) ɚ

0.85 (1.22)

Mean diameter of midshaft (TAD)

24/16

1.74 (0.75)

0.82 (0.94)

0.97 (0.83) ɚ

Minimal diameter of midshaft (TD min)

27/18

1.43 (0.66)

1.51 (1.82) b

1.43 (1.13) ɚ

Maximal diameter of midshaft (TD max)

27/15

0.00 (–0.44)

–2.20 (–1.59) b

–0.86 (–0.85) ɚ

T10. Midshaft circumference

27/17

0.56 (0.10)

0.00 (0.10)

0.27 (0.10)

Note: n1 and n2 are numbers of observations in male and female samples, respectively; signi¿cance level of differences between the sides: a – p < 0.05, b – p < 0.01, c – p < 0.001; scores with a signi¿cantly non-normal distribution are italicized; those indicating signi¿cant sexual differences are set in bold; those showing signi¿cant temporal differences are underscored.

Table 3. Directional asymmetry indexes of clavicles and limb bones in the Staroaleyka sample* Trait

n1/n2

1

2

%DA Males

Females

Pooled

3

4

5

Clavicula C1. Length

12/8

–1.01 (–1.27) ɚ

–2.06 (–1.28)

–1.19 (–1.27) b

C4. Vertical diameter

12/10

1.04 (4.63)

2.11 (2.59)

1.57 (3.70) ɚ

C5. Sagittal diameter

12/10

–0.27 (–2.06)

1.84 (3.11)

1.32 (0.29)

b

1.74 (1.85) 1.57 (1.26)

Mean diameter (CAD)

12/10

0.96 (0.88)

2.78 (3.01)

C6. Circumference

11/10

0.00 (–0.50)

2.86 (3.21) b

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Table 3 (continued) 1

2

3

4

5

Humerus H1. Maximal length

14/20

1.38 (1.32) b

1.80 (1.62) c

1.58 (1.50) c

H3. Upper epiphysis breadth

12/17

1.43 (1.30) ɚ

1.21 (1.48) b

1.21 (1.41) c

H4. Lower epiphysis breadth

11/14

0.00 (–0.37)

–0.42 (–0.09)

0.00 (–0.22)

H5. Maximal diameter of midshaft

15/19

0.40 (0.13)

0.53 (1.60)

0.43 (0.95)

H6. Minimal diameter of midshaft

15/20

–0.49 (–0.82)

0.33 (1.33)

0.00 (0.41)

H6b. Midshaft breadth

15/20

–0.42 (–0.22)

0.27 (0.73)

0.00 (0.27)

3.86 (3.01)

2.60 (2.95) c

b

H6c. Sagittal diameter of midshaft

15/19

1.63 (2.72)

Mean diameter of midshaft (HAD)

15/19

1.04 (1.20)

2.49 (1.87)

2.11 (1.58) b

Midshaft breadth at 35 % length (H35 % ml)

15/20

1.06 (0.36)

–0.61 (0.16)

0.00 (0.24)

Sagittal diameter at 35 % length (H35 % ap)

15/20

0.00 (–0.01)

1.82 (1.89)

1.05 (1.07) ɚ

Mean diameter at 35 % length (HAD35 %)

15/20

0.00 (0.19)

1.29 (1.05)

1.20 (0.68)

H7. Minimal circumference of midshaft

15/20

0.00 (0.03)

0.87 (0.99)

0.00 (0.58)

ɚ

ɚ

b

1.54 (1.24) b

H7a. Circumference of midshaft

15/19

0.00 (0.47)

1.72 (1.84)

H10. Vertical diameter of head

12/19

–0.42 (0.07)

1.50 (1.05) ɚ

0.47 (0.67)

Radius R1. Maximal length

12/18

0.42 (0.47) ɚ

1.03 (1.15) c

0.62 (0.88) c

R4. Maximal breadth of midshaft

15/19

2.25 (3.21) b

2.48 (1.82) ɚ

2.46 (2.44) ɚ

R5. Sagittal diameter of shaft at the maximal breadth level

15/19

0.79 (–0.12)

0.99 (0.80)

0.93 (0.39)

Mean diameter of shaft at the maximal breadth level (RAD max)

15/19

1.39 (1.79)

1.83 (1.43)

1.55 (1.59) ɚ

R4a. Midshaft breadth

12/18

2.32 (3.00) ɚ

5.33 (5.19) c

4.52 (4.44) c

R5a. Sagittal diameter of midshaft

12/18

–0.36 (–0.28)

0.00 (0.85)

0.00 (0.22)

Mean diameter of midshaft (RAD)

12/18

1.50 (1.60) ɚ

2.81 (3.30) c

2.06 (2.59) c

R3. Minimal circumference of midshaft

14/20

0.00 (–0.35)

1.36 (1.02)

0.54 (0.45)

Distal epiphysis breadth (R dist)

12/13

0.00 (–0.17)

2.33 (1.60)

0.71 (0.75)

ɚ

Ulna U1. Maximal length

11/15

0.19 (0.55)

1.32 (1.27) c

1.10 (0.97) c

U11. Sagittal diameter at maximal breadth level

16/20

–1.41 (0.19)

0.00 (0.07)

0.00 (0.12)

U12. Maximal midshaft breadth

16/20

–1.79 (–0.20)

1.25 (2.68) ɚ

0.81 (1.40)

Mean diameter at the maximal breadth level (UAD max)

16/20

–0.56 (–0.05)

0.43 (1.54)

0.15 (0.84)

U11a. Sagittal diameter of midshaft

12/16

–2.12 (–2.40)

–1.71 (0.23)

–1.71 (–0.90)

U12a. Midshaft breadth

11/16

–2.43 (–3.47)

ɚ

1.15 (1.28)

–0.62 (–0.76)

Mean diameter of midshaft (UAD)

12/13

–2.73 (–2.95)

ɚ

1.26 (0.85)

–1.12 (0.78)

U3. Minimal shaft circumference

13/20

0.0 (0.33)

1.47 (0.71)

1.29 (0.56)

Femur F1. Maximal length

15/21

–0.64 (–0.25)

–0.50 (–0.44) ɚ

–0.57 (–0.36) ɚ

F6. Sagittal diameter of midshaft

15/23

1.45 (1.31) ɚ

–0.44 (–0.03)

0.00 (0.40)

b

–2.45 (–3.21) c

F7a. Transverse diameter at midshaft

16/23

–1.20 (–1.70)

–4.46 (–2.95)

Mean diameter of midshaft (FAD)

15/23

0.00 (–0.09)

–2.02 (–1.56) b

–1.04 (–1.06) b

F8. Midshaft circumference

15/22

0.00 (–0.18)

–1.64 (–1.43) b

–0.67 (–0.92) b

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147

Table 3 (end) 1 F21. Lower epiphysis breadth F18. Caput height

2

3

4

5

14/17

0.60 (0.31)

0.00 (–0.08)

0.00 (0.09)

14/15

–0.44 (0.25)

1.11 (0.67)

0.72 (0.47)

Tibia T1. Full length

16/22

–0.29 (–0.10)

–0.30 (–0.14)

–0.30 (–0.12)

T5. Maximal breadth of upper epiphysis

9/13

–0.66 (–0.43)

0.00 (0.07)

0.00 (–0.14)

T6. Maximal breadth of lower epiphysis

15/16

0.95 (1.07)

0.52 (0.71)

0.95 (0.88) ɚ

T8. Sagittal diameter of midshaft

16/22

0.20 (–0.17)

–0.66 (–1.81)

–0.65 (0.00)

T8a. Sagittal diameter of shaft at the level of nutrient foramen

16/22

0.96 (1.05)

–0.38 (–0.31)

0.45 (0.93) 1.81 (1.02)

T9. Transverse diameter of midshaft

15/23

4.32 (1.44)

1.39 (2.62)

T9a. Transverse diameter of shaft at the level of nutrient foramen

16/23

1.58 (1.72)

1.09 (0.88)

1.09 (1.09) ɚ

Mean diameter of midshaft (TAD)

15/22

0.00 (0.51)

0.48 (0.21)

0.51 (0.48)

Minimal diameter of midshaft (TD min)

17/23

0.00 (–0.50)

2.17 (2.43) b

0.57 (1.19) ɚ

Maximan diameter of midshaft (TD max)

17/23

0.94 (0.21)

–1.54 (–1.16)

–1.10 (–0.58)

T10. Midshaft circumference

16/22

0.00 (0.24)

–0.68 (–0.15)

0.00 (0.02)

ɚ

*See Note to Table 2. Fig. 1. Variation ranges of %DA of humeral length (H1) and ulnar length (U1). Chronological differences. a – males; b – females, a, b – 25–75 %; c – median.

more asymmetric vertical diameter of humeral heads H10 (ɪ = 0.041) (Fig. 2). Diachronic changes are evident in the asymmetry of the circumference (ɪ = 0.003), of the vertical (ɪ = 0.035), and midshaft (ɪ = 0.037) diameters of the clavicle and of the midshaft width of the humerus H6b in females (ɪ = 0.044) as well as in the width of the lower epiphysis H4 in male (ɪ = 0.025), female (ɪ = 0.003) and pooled (ɪ = 0.000) groups (Fig. 3). Sex differences in %DA scores of ulnae are seen in the AC sample: R4a shaft width and R5a : R4a ratio are less asymmetric in females

ɚ

c

b

Table 4. Proportions of individuals with right and left bias of humeral length (ɇ1), % Sample

Sex

n

Right-bias

” 0.5

Left-bias

AC

ƃ

19

57.9

26.3

15.8

Ƃ

21

57.1

19.0

23.8

ƃ+Ƃ

40

57.5

22.5

20.0

ƃ

14

85.7

7.1

7.1

Ƃ

20

80.0

20.0

0.0

ƃ+Ƃ

34

82.4

14.7

2.9

SC

Note: Proportions showing signi¿cant differences between samples (ɪ < 0.05) are italicized.

148

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Table 5. Mean shaft cross-section index scores Index

Andronovo

Trait code

Humeral midshaft cross-section Same Same, at 35 % level

ƃ

Staroaleika Ƃ

H6 : H5

76.3/77.8 (21)

H6ɫ : H6b

97.5/95.0 (21)

ɚ

ƃ

72.5/75.1 (23)

b

101.5/95.2 (22)

c

Ƃ

76.8/77.5 (15)

74.4/74.7 (19)

97.5/94.8 (15)

93.1 /91.1 (19)

ɚ

H35 % ap : H35 % ml 109.1/108.6 (18) 113.1/113.9 (17) 106.5/106.8 (15) 103.8/101.9 (20)

Radial midshaft cross-section

R5 : R4

Same

69.8/74.6 (16) ɚ c

68.4/70.1 (14)

70.1/72.8 (15) ɚ

68.2/68.9 (19)

73.9/77.4 (15)

78.2/80.8 (12)

ɚ

73.4/76.8 (18) b

R5a : R4a

72.5/82.3 (15)

Pilastry

F6 : F7ɚ

101.1/99.3 (27) ɚ

98.5/96.0 (22)

101.0/98.1 (15)

94.9/92.3 (23) ɚ

Tibial midshaft cross-section

Ɍ9 : Ɍ8

86.1/84.0 (24)

89.2/83.6 (16) b

86.9/85.2 (15)

85.5/81.8 (22) ɚ

Ɍ9ɚ : Ɍ8ɚ

75.9/74.5 (21)

80.7/75.9 (17) b

77.5/76.9 (16)

75.9/74.8 (22)

Cnemia

ɚ

Note: Indexes refer to right/left sides; number of observations is indicated in parentheses; signi¿cance of differences in %DA scores: a – ɪ < 0.05, b – p < 0.01, c – p < 0.001; %DA scores showing signi¿cant sexual differences are italicized; those showing signi¿cant temporal differences are set in bold.

Table 6. Coef¿cients of correlation between the DA of upper limb bone lengths and breadths Traits

Andronovo ƃ

Staroaleika ƃ+Ƃ

Ƃ

Pooled

ƃ

Ƃ

ƃ+Ƃ

ƃ+Ƃ

H1 H4

0.240 (15)

0.668 (14)

0.422 (29)

–0.394 (11)

0.506 (13)

0.177 (24)

0.089 (53)

HAD

0.085 (18)

0.410 (20)

0.300 (38)

0.538 (14)

0.386 (19)

0.424 (33)

0.300 (71)

R1

0.028 (12)

0.466 (7)

0.146 (19)

0.399 (8)

0.422 (17)

0.400 (25)

0.296 (44)

HAD

0.305 (13)

0.612 (13)

0.533 (26)

–0.003 (8)

–0.055 (17)

–0.120 (25)

0.321 (51)

R1

–0.018 (14)

0.692 (9)

0.193 (23)

–0.552 (12)

0.282 (18)

0.156 (30)

0.172 (53)

RAD

Note: Observation number is indicated in parentheses; values signi¿cant at the 0.05 level are set in bold.

ɚ

b

c

d

e

Fig. 2. Variation ranges of %DA of the transverse dimensions of clavicles (C) and humeri (H). Sexual differences. a – males; b – females, a, b – 25–75 %; c – median; d – variation range without extreme values; e – extreme values.

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(p = 0.041 and 0.009, respectively). Male groups differ in the asymmetry of midshaft cross-section asymmetry of radii: the %DA scores of R4a are higher in the earlier group (ɪ = 0.002), and the same applies to R5a (ɪ = 0.037) and R5a : R4a (p = 0.000) (Fig. 4). Leg bones. The left femora and tibiae are longer than the right ones. Femoral midshaft width F7a displays leftside asymmetry, which is more pronounced in females. The sagittal midshaft diameter F6 in Early Iron Age males shows a signi¿cant right-bias. Sex differences in the %DA of the femoral midshaft in the SC group are close to the signi¿cance level (ɪ = 0.09). Bronze Age males and Early Iron Age females show signi¿cant asymmetry of the pilastry index, whereas Bronze Age males and Early Iron Age females reveal signi¿cant asymmetry in the pilastry index F6 : F7a. No directional asymmetry in the width of the lower epiphysis of femora was detected.

Medio-lateral and minimal shaft diameters of the tibiae show a right-bias, whereas the sagittal and maximal diameters display a predominance of the left side. These tendencies are more expressed in females, especially in the AC group. In female samples, cross-section shaft ratios T9 : T8 and T9a : T8a are signi¿cantly asymmetric too. Most %DA scores relating to the epiphyses of tibiae are close to zero. Sex differences are observed in %DA of the maximal midshaft diameter of tibiae (TD max) in the AC sample (ɪ = 0.049) and of the minimal diameter (TD min) in the SC sample (ɪ = 0.011). Diachronic differences are seen in %DA of Ɍ8ɚ (sexes pooled, ɪ = 0.017) and of the cnemic index Ɍ9a : Ɍ8a in females (ɪ = 0.034) (Fig. 5). To assess the %DA scores in skeletal series of pastoralists from the Altai, the between-group scale was used. Vertical lines in Fig. 6 and 7 show variation limits of median %DA scores in geographically and

Fig. 3. Variation ranges of %DA of the transverse dimensions of clavicles (C) and humeri (H). Chronological differences. See Fig. 2 for explanations.

Fig. 4. Variation ranges of %DA of radial (R) and ulnar (U) shafts. Sexual and chronological differences. See Fig. 2 for explanations.

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150

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Fig. 5. Variation ranges of %DA of tibial shaft (Ɍ). Sexual and chronological differences. See Fig. 2 for explanations.

Fig. 6. %DA of clavicles and arm bones. Between-group scale.

Fig. 7. %DA of leg bones. Between-group scale.

chronologically diverse groups (Auerbach, Ruff, 2006; Auerbach, Raxter, 2008) and the position of samples from the Altai. The between-group variation ranges of %DA scores of transverse dimensions of shafts are much higher than those of the longitudinal dimensions. Arm bones and clavicles in the SC group reveal relatively high %DA scores, the radius in males being an exception. In the AC series, however, the %DA scores are low. If the 24 groups used for comparison are arranged in the increasing order of %DA of humeri, then the male AC group surpasses seven of them, and

the female AC group surpasses six. The SC samples, by contrast, surpass 19 and 17 series, respectively. The asymmetry of the transverse shaft diameter of humeri is quite low in males and below average in females. In that characteristic, the female AC series surpasses three groups out of 24 and the SC series, nine. In terms of the %DA of the ulnar midshaft, AC males surpass seven groups out of nine, and AC females, four; the respective numbers for the SC sample are three and six. A strong left-bias of the clavicular shaft width in the female AC sample may be due to its small size. The absolute

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151

sex dimorphism of %DA of transverse dimensions of humeral shafts is moderate (Fig. 8), and that of radial shafts, high (Fig. 9). The %DA of leg bone lengths is close to zero or slightly negative. In both AC and SC female samples, an extremely high left-bias of the transverse dimensions of femoral shafts is observed. The distinctive feature of the Altai groups is that the %DA of the transverse dimensions of tibial shafts is very small or positive (Fig. 7). Discussion The DA of long bones mirrors functional asymmetries of locomotor behavior. Dominant limbs surpass their counterparts in strength, precision, speed, and coordination of movements, whereas the role of nondominant limbs is auxiliary. Most modern people (85–90 %, according to various estimates) are right-handers (Porac, Coren, 1981; Annett, 1985; Bragina, Dobrokhotova, 1988). Bones of the dominant arm are longer and thicker. The share of individuals showing right-bias in the longitudinal dimensions of humeri (or humeri plus radii) in ancient populations is often close to the proportion of righthanders in modern groups (Schultz, 1937; Steele, Mays, 1995; ýuk, Leben-Seljak, Štefanþiþ, 2001; Auerbach, Ruff, 2006). Directional asymmetry of the clavicle, too, is related to the functional dominance of the right arm (Auerbach, Raxter, 2008). The functional asymmetry of the organism, mirrored by the directional asymmetry of limb bones, is known to be genetically determined. However, populations with the same share of right-handers may differ in morphological asymmetry depending on the nature and amount of habitual physical exercise. The role of mechanical loading in the morphological DA of arm bones has been studied in high-ranking athletes (Jones et al., 1977; Krahl et al., 1994; Haapasalo et al., 1996, 2000; Kontulainen et al., 2003; Shaw, Stock, 2009a; and others) as well as in people not engaged in vigorous physical activity (Blackburn, Knüsel, 2006; Özener, 2007, 2010). The study of geographically diverse skeletal series reveals considerable variation of the DA of structural components of bones such as length, width of shafts and articular surfaces. Diaphyseal breadth exhibits the most pronounced asymmetry, bone length is the least asymmetric, and epiphyses, speci¿cally articular and periarticular surfaces, are intermediate (Trinkaus, Churchill, Ruff, 1994; Auerbach, Ruff, 2006). No correlation between the asymmetry of bone length and that of shaft width was found (Auerbach, Ruff, 2006). Considerable between-group and within-group variation in the asymmetry of diaphyseal breadth testi¿es to the plasticity of this trait (Auerbach, Ruff, 2006; Auerbach, 2007; Auerbach, Raxter, 2008). Longitudinal growth

Fig. 8. Sexual dimorphism of %DA of the midshaft diameter of humeri. Difference between male and female scores. Groups: 1 – AC; 2 – SC; 3–11 – after (Auerbach, Ruff, 2006); 12–26 – after (Auerbach, Raxter, 2008).

Fig. 9. Sexual dimorphism of %DA of the midshaft of the radius. Difference between male and female scores. Groups: 1 – AC; 2 – SC; 3–11 – after (Auerbach, Ruff, 2006).

of bones, by contrast, is more influenced by heredity (Hallgrímsson, Willmore, Haall, 2002). The negative relationship between the canalization of bone dimensions and the effect of mechanical stimuli in DA is evident during development. In children below one year of age the humeri are normally symmetric, whereas the asymmetry is Àuctuating and concerns disphyseal and epiphyseal width rather than the length of bones. Directional asymmetry in humeral morphology develops parallel to functional asymmetry. The right-side asymmetry of diaphyseal breadth progresses at the highest rate, whereas that of humeral length develops with the slowest rate (Blackburn, 2004). In sum, the decrease or increase in the asymmetry of diaphyseal breadth mirrors both between-group and

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within-group differences in the nature and magnitude of physical stress. Asymmetry in upper limb bones increases if the “division of labor” exists between the arms. Conversely, asymmetry decreases if the functional differences between the arms are minor. Lateralization of manual effort also depends on the complexity of operations. During simple movements requiring mostly physical force, the functional difference between the dominant and nondominant arm is lower than in tasks which demand precision movements, coordination, and visual control, whereby the dominant hand is normally employed (Bryden, 2002). Published data suggest that the %DA of diaphyseal breadth of upper limb bones can reflect changes in economy and lifestyle, speci¿cally in sedentism versus nomadism. In males, %DA of humeri is higher in the least sedentary groups (Auerbach, 2007). In most agriculturalists, humeral and clavicular shafts are less asymmetric than in hunter-gatherers (Auerbach, Raxter, 2008). In the male pastoralist samples from the Altai, %DA scores of humeral and clavicular shaft width are very low; they are higher in female samples, especially in that from SC burials. As the analysis of humeral strength indicates, physical loading was high in both groups of males and in AC females while being low in SC females (Tur, 2013). In males, therefore, the loading was bilateral and involved mostly physical strength, whereas the work of females was more related to precision, and the dominant hand was evidently used more often. Sexual differences in the asymmetry of mechanical loading effects indicates gender division of labor among the pastoralists of the Altai. Decrease of such effects on arm bones in the Early Iron Age, accompanied by a higher role of the dominant hand, may be caused by the technological sophistication of female labor. The opposed dominance of transverse shaft dimensions of radii and ulnae, observed in males of both groups, probably testifies to an unequal distribution of stress between forearm bones. Bronze Age and Iron Age males differ mainly in the %DA of size and especially shape of the cross-section of radial shafts. This may be because the mechanisms of adaptation to physical stress are not the same in the proximal and distal segments of arm bones. Changes in the cross-section shape of the humeral shaft correlate with the increase in circumference; changes in the shape of the shaft cross-section secure an optimal energetic balance between bone strength and bone weight (Stock, 2006; Shaw, Stock, 2009a). Humeri reÀect the effects of stress more adequately than forearm bones do (Ibid.). Difference in the DA of the shaft cross-section shape of radii and of the lower epiphysis width of humeri in male AC and SC samples suggests that the nature of physical stress on arm bones in males changed in the Early Iron Age.

The DA variation of upper bone length is apparently caused by several factors. The %DA of humeral, radial, and ulnar length is higher in females than in males, but the differences are insigni¿cant possibly because the sample size is small. Greater right-bias of arm bone length in females is frequent in other groups as well (Auerbach, Ruff, 2006; Auerbach, Raxter, 2008), although there are exceptions (Steele, Mays, 1995; Papaloucas et al., 2008). This is due to the normally higher share of left-handers among males (Bryden, 1982; McManus, 1991; Porac, Coren, 1981), and this parameter varies at the population level. The DA of arm bone length in the pastoralists of the Altai shows signi¿cant diachronic changes. Both male and female Iron Age samples display a higher %DA of humeral length, and the same is true of forearm bones in females. The share of individuals with the right-bias of humeral length in the SC group is roughly the same as that of right-handers in the modern population, whereas in the AC group it is much lower. Diachronic changes of the DA of arm bone length may be caused by mechanical loading, as evidenced by certain statistically signi¿cant correlations between the %DA of humeral length and transverse dimensions (Table 6). The effect of high physical stress on the length of bones of the dominant arm has been observed in highranking tennis players (Krahl et al., 1994), although such facts are rare. The study of age changes of the DA of arm bones reveals signi¿cant variation in the asymmetry level during puberty (11–14 years of age), indicating heterogeneity of effects (Chermit, Aganyants, 2006) and suggesting that high heritability notwithstanding, bone length becomes more sensitive to mechanic stress during the pubertal period. Absence of correlation between the asymmetry of bone length and that of shaft width in a large sample from various geographic regions (Auerbach, Ruff, 2006) can be explained by the heterogeneity of the sample and by opposed tendencies in various local groups. Apparently, physical stress during puberty is not always high enough to affect bone length. Apart from the mechanical factor, DA of arm bone length can be affected by environmental and genetic stress. The neural substrate for right-handedness is formed at the early stages of intrauterine development. As the results of the ultrasound study demonstrate, 92 % of fetuses suck their right thumbs (Hepper, Shahidullah, White, 1991). The shift of the genetically determined level of functional lateralization of the hands, speci¿cally the increase of the left-side dominance, may be due to the destabilization of the intrauterine development caused by homozygosity in many loci, by the unfavorable factors such as infection and, especially, by the combination of such factors (Markow, 1992; Yeo, Gangestad, 1993). Available data suggest that the frequency of left-handers in certain Arctic populations may be as high as 33.8 % (Stepanov, 1988).

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Environmental stress can apparently affect the DA of arm bone length during skeletal maturation. While the fusion of the epiphyses with the metaphyses normally proceeds synchronically on both sides, certain skeletal populations with a high occurrence of episodic and cumulative stress markers display signi¿cant DA and FA in the synostosis of the humeri. Early synostosis is more often observed on the right side (Albert, Greene, 1999). The lesser lateralization of arm bone length in the pastoralists of the Altai, then, can be caused by environmental and/or genetic stress. To test this idea, one will need to focus on traits with an “ideal” FA. Functional asymmetry of the lower limbs is less evident than that of the upper ones. The dominant leg is more often used for movements involving motor coordination, whereas the other leg is used for support and for maintaining balance (Gabbard, Hart, 1996; Sadeghi et al., 2000). In most right-handers, the dominant leg is the right one (Gabbard, Iteya, 1996). The direction of functional and morphological asymmetry of the legs differs from that of the arms. Bones of the left leg, normally used for support, are larger. This contralateral, or “crossed” symmetry is due to the fact that the non-dominant leg is subjected to a heavier stress under the weight of the body. Published data suggest that the femoral length and shaft breadth show a signi¿cant leftbias. Tibial length, however, displays no asymmetry, whereas tibial shaft breadth is greater on the left side in some groups (Auerbach, Ruff, 2006) and on the right side in others (Auerbach, 2007). The DA of leg bones is weaker than that of arm bones. Apparently locomotion involves more symmetric stress. In addition, it limits the asymmetry of leg bone length. Cross-section shape and strength of long bone shafts correlate with the mobility level (Ruff, 1987; Stock, Pfeiffer, 2001; Stock, 2006; Shaw, Stock, 2009b). Female AC and SC samples, unlike male ones, show very high negative scores on %DA of humeral shaft breadth. The left-bias of the mean femoral shaft diameter is normally less than 1 % (Auerbach, Ruff, 2006). However, in some female groups such as pre-industrial Europeans (Ibid.) and American Indians who practiced both foraging and farming (Wescott, Cunningham, 2006; Auerbach, 2007), the scores are higher. Because mobility level affects the sagittal shaft diameter of humeri more than the medio-lateral diameter (Ruff, 1987; Stock, Pfeiffer, 2001; Stock, 2006), a higher left-bias of the latter may be caused by speci¿c features of physical activity under a sedentary lifestyle. According to the world summary published by Auerbach and Ruff (2006), the tibial shaft breadth displays a left-bias. This tendency, however, is not universal (Auerbach, 2007), and our data provide yet another exception. This lack of uniformity may be due

to the fact that the morphological adaptation of the distal segments of leg bones, as well as that of arm bones, is mostly related to shape changes (Stock, 2006). The DA of size and shape of the cross-section of tibial shafts in AC and SC groups shows sexual and chronological differences. In females, the left tibiae have a lesser mediolateral (minimal) diameter and/or a larger sagittal (maximal) diameter, resulting in greater lateral Àattening (platycnemia). The same tendency has been observed in certain other groups (Ruff, Jones, 1981; Bagashev, 1993; Wanner et al., 2007), but the underlying factors remain poorly understood. Data relating to the asymmetry of stress on the legs during walking and running with regard to different functional specialization of the right and the left leg (mobility versus stability) are contradictory (Sadeghi et al., 2000; Seeley, Umberger, Shapiro, 2008). Factors unrelated to physical exercise may affect the DA of leg bones as well. Conclusions The analysis of the DA of long bones in geographically and chronologically diverse populations contributes to the understanding of the functional adaptation of the skeleton and provides data for the comparative study of physical activity with regard to lifestyle and the division of labor between the sexes. Results relating to the Bronze Age and Early Iron Age pastoralists of the forest-steppe Altai uphold the existing view that the DA of the shaft breadth is more variable than that of bone length. We have also demonstrated that the DA of the proximal segments of limbs relates to both size and shape of the shaft cross-section whereas in the distal segments mostly the shape is affected. The DA of the longitudinal and transverse dimensions of arm bones are correlated. Tibial shaft breadth shows a right-bias. The DA of transverse dimensions of arm bones and clavicles displays both sexual and chronological differences. Results of the analysis of the DA of upper limb shaft breadth suggest that male pastoralists experienced bilateral manual stress mostly involving physical effort. Female labor required more accurate movements, and the dominant arm was engaged more often. This tendency became more prominent in the Early Iron Age. At the same time, women experienced more asymmetric stress on leg bones. The DA of arm bone length was low in the Bronze Age but high in the Early Iron Age. Chronological differences in the asymmetry of these dimensions may be affected by the mechanical factor as well as by the environmental and/or genetic stress. To specify the relative role of these factors, new data on FA are needed.

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