Professional Documents
Culture Documents
William L. Hylander
Department of Biological Anthropology and Anatomy and Primate Center, Duke University, Durham, NC 27710, U.S.A.
Can a bimanual activity such as thrusting a spear during hunting produce bilateral asymmetries in the strength of the
upper limbs? This question is important to arguments about the predatory capabilities of Neandertals and early modern
humans. To address this question, we determined the magnitude and direction of reaction forces on the upper limbs
during thrusting spear use. We collected lateral video records of eight adults thrusting an instrumented aluminum rod
into a padded target. This ‘‘spear’’ was instrumented with two sets of four strain gauges placed at two positions along
the shaft to register the force along the shaft and the distribution of those forces relative to the two limbs. From the
gauge output and video we were able to calculate loads experienced by the trailing limb (holding the proximal spear)
and the leading limb (holding the distal spear) as well as approximate bending moments along the trailing limb. The
trailing limb provides a significantly greater portion of the force during spear impact and when the spear is held
forcefully on the target. The loads on this limb at spear impact are twice body weight and the bending moments on the
trailing humerus are large and appear to occur primarily in the parasagittal plane. These data, in combination with
fossil humeral cross-sectional data and the lack of evidence for throwing spears among Eurasian Neandertals, suggest
that previously documented humeral strength asymmetries in Eurasian Neandertals and early Upper Paleolithic
Modern human males can be plausibly linked to spear thrusting. 2002 Elsevier Science Ltd. All rights reserved.
1
Based on cross-sectional properties of the humeral diaphysis as measured at 35% of length from the distal end: All
data from Churchill et al., 1996.
2
CA: cortical area (mm2: a measure of the thickness of the diaphyseal cortical bone and resistance to compressive
and tensile loads); J: polar moment of inertia (mm4: a measure of diaphyseal rigidity to bending and torsional
moments). Percent asymmetry calculated as [(max–min)/min]*100. Median, quartile range, range, and sample n
provided.
3
Ratio of the second moment of inertia (I; mm4; a measure of diaphyseal rigidity to bending moments in a single
plane) in the anteroposterior plane (Ix) to that of the mediolateral plane (Iy). Mean, SD, sample n provided.
4
Ratio calculated using mean Ix and Iy values: data from Fresia et al., 1990.
the Middle and early Upper Paleolithic of western humeral shafts that are rounder in section, and hence
Eurasia? that were roughly equally resistant to bending
moments in both directions. The rounded shaft cross-
sections seen in the humeri of the late Upper Paleo-
lithic sample are consistent with resistance to torsional
Background loads as produced during throwing (Tullos & King,
Consideration of humeral diaphyseal cross-sectional 1973; Gainor et al., 1980; Pappas, Zawacki & Sullivan,
shape in Neandertals and early modern Europeans led 1985). The greater anteroposterior bending strength of
Churchill and colleagues (1996) to argue against the Neandertal and early Upper Paleolithic male
throwing as the source of the mechanical loads respon- humeri has been argued to reflect resistance to large
sible for the observed asymmetry. They considered the bending moments engendered by thrusting spear use
implications of size asymmetry in Neandertals and (Churchill, Weaver & Niewoehner, 1996). Further-
early Upper Paleolithic modern humans and the more, Churchill et al. suggested that forceful employ-
shape differences between this group and late Upper ment of a thrusting spear, although seemingly
Paleolithic humans. bimanual in nature, may in fact generate greater reac-
Neandertal and early Upper Paleolithic (Aurigna- tion forces and larger bending moments in the ’’trail-
cian and Gravettian) modern human males are ing’’ limb (the limb holding the proximal end of the
characterized by humeral shafts that are wider antero- spear) than in the ‘‘leading’’ limb (on the distal spear).
posteriorly than mediolaterally. In addition, the right General observations suggest that the dominant limb
humeri in these specimens are considerably more (the right-hand in most people) is usually employed as
robust, and therefore are likely to be stronger in all the trailing limb. Thus, they argued, that regular
directions compared to left humeri (Table 1) close-range hunting with thrusting spears may have
(Churchill, Weaver & Niewoehner, 1996). Late Upper produced the pattern of right-dominant humeral
Paleolithic males, on the other hand, tend to have strength asymmetry seen in these groups.
Spear Use in Pleistocene Homo 105
We provide here the results of experimental research field javelins]), Thieme (1997, 1999) has argued that
aimed at testing the claims that (1) bimanual spear these spears were designed for aerodynamic stability
thrusting produces asymmetrical loads in the two limbs and hence projectile use, and that they were used to kill
(and thus could be the basis for differential modelling horses, as represented by the remains of at least 15
responses in the two humeri and ultimately, bilateral animals in this level. Similar artifacts have been recov-
strength asymmetry) and (2) thrusting spear use ered from Hoxnian Interglacial (OIS 13) aged deposits
engenders relatively large anteroposterior bending at Clacton-on-Sea, England (the distal 39 cm of a
moments on the humerus of the dominant (trailing) sharpened Yew stave: Warren, 1911; Oakley et al.,
limb. 1977) and from Eemian Interglacial (OIS 5) deposits at
Lehringen, Germany (a 2·4 m long, broken spear of
Yew, found between the ribs of a elephant skeleton:
Close range weapons of the Middle and Early Upper Movius, 1950; Tode, 1954).
Paleolithic There is lithic-use wear evidence that during Middle
The earliest direct evidence of long range projectile Paleolithic times humans began to haft lithic points to
weaponry comes in the form of the remains of spear- spears or handles to make composite weapons (Shea,
throwers (atlatls or propulseurs) from Solutrean con- 1988, 1989; Beyries, 1990; Shea et al., 2001). The
texts at La Placard (Breuil, 1913) and Combe-Saunière recovery of a broken Levallois point embedded in a
(Cattelain, 1989), dating to c. 19–17,000 years ago. cervical vertebra of a wild ass at Umm El Tlel (Boëda
There is presently only indirect evidence suggesting et al., 1999) further testifies to the use of lithic weapon
that the spear-thrower (or other forms of long range armatures in the Mousterian. However, the extent to
projectiles) may have come into common use earlier. which this innovation reflected a concern for projectile
From the late Middle Paleolithic through the Early penetration versus the overall effectiveness (or reliabil-
Upper Paleolithic certain design features occur with ity sensu Bleed, 1986) of a hand-delivered weapon
increasing regularity in lithic (and in some cases is not clear (see Shea, 1997). Edgewear damage on
osseous: Gaudzinski, 1999) points. These include Mousterian and Levallois points also supports the
relatively small size, symmetry about the long axis, notion that they were sometimes used as weapon
basal modification to facilitate hafting (including the armatures (Shea, 1989, 1990, 1997), but is insufficient
use of stems and shoulders), and size standardization for differentiating projectile (javelin) versus hand
of the proximal end—traits that can be variously thrust (lance) modes of deployment (Shea, 1993). Thus,
seen in points that predate the earliest known while analysis of lithic armatures may better elucidate
atlatls—including Aterian, Teyjat, Font Robert, the characteristics of the hunting weapons themselves,
El-Wad (Font Yves) and Gravette points (see Straus, they do not speak to the question of how these
1990a; Peterkin, 1993). These design elements may weapons were deployed. Shea, Davis & Brown (2001)
reflect a concern for projectile aerodynamics and pen- has recently shown the effectiveness of Levallois
etrating capabilities (see Guthrie, 1983; Christenson, points of the Levantine Mousterian in piercing goat
1986; Odell & Cowan, 1986; Shea, Davis & Brown, carcasses in simulated calibrated thrusting conditions.
2001), and hence their appearance in the Pleniglacial Shea, Davis & Brown (2001) noted in particular that
may herald the advent of real projectile weaponry. the lack of data on thrusting forces hinders analyses of
Thus sometime during the late Middle Paleolithic, the use of spears by Neandertals and early modern
early Upper Paleolithic or earliest late Upper Paleo- humans.
lithic, hunters increasingly employed long range While some of the recovered fossil spears exhibit
projectile weaponry. Prior to this hunters were design elements that may reflect a concern for aerody-
probably equipped with hand-held, close range namics, their overall size and shape is most similar to
hunting technology, which likely included an arsenal of that of ethnographically known thrusting spears (Table
sticks, clubs and stones (see e.g., Thieme, 1999), 2) (Oakley et al., 1977). The Schöningen and Lehringen
but which undoubtedly emphasized hand-delivered, spears are relatively thick compared to recent throwing
as opposed to spear-thrower-delivered (i.e. atlatl), spears (Table 2), as is the Clacton spear (the maximum
spears. diameter of the preserved portion of the distal end is
Large, heavy pointed staves have been recovered 39 mm: Oakley et al., 1977). Indeed, the average maxi-
from a number of Middle and early Late Pleistocene mum diameter of four of the fossil spears (for which
sites in Europe. The most remarkable of these was the published data are available) is almost 3·5 standard
discovery, between 1995 and 1997, of seven largely deviations above the mean for eight recent thrusting
intact wooden spears from the c. 400,000 year old (OIS spears and is more than seven standard deviations
11) level 4 of Schöningen 13, near Helmstedt, Germany above the mean for 28 modern throwing spears
(Thieme, 1997, 1999). These spears range in length reported by Oakley et al. (1977) (Table 2). When
from 1·8–2·5 m and in maximum thickness from 29– maximum diameter is taken relative to spear length
47 mm (Thieme, 1999). Based on design attributes (thickness index: Table 2), the fossil spears fall between
(namely, distal positioning of the centre of mass and ethnographically known thrusting spears and digging
tapering of the proximal end [as with modern track and sticks. While the published data on the mass of the
106 D. Schmitt et al.
Maximum
Length Mass diameter Thickness
(mm) (g) (mm) index1 Source
1
100*Maximum diameter/length.
fossil specimens is not available, the great length and Behavioural inferences from structural analysis of
width of the fossil spears suggests that they would have human fossil remains
been heavy relative even to modern thrusting spears,
and easily four to five times the weight of modern It bears mentioning why we expect either of these
throwing spears (Table 2). Clearly, if these were behaviours (throwing or thrusting) to be reflected in
designed to be projectile weapons, they were larger the skeletal morphology of the hominids themselves.
and heavier than any comparable projectile used by As mentioned, living bone tissue is known to adap-
modern foragers. tively model and remodel in response to biomechanical
We should note at the outset that, with respect to stimuli (see references in Ruff, 1992; Trinkaus et al.,
hand-delivered spears, the distinction between thrust- 1994). Thus variation in the relative amount and
ing and throwing spears is largely artificial: recent distribution of cortical tissue in long bone diaphyses is
(historically known and extant) hunter-gatherers use argued to reflect interindividual variation in the inten-
hand-held spears in both manners (Churchill, 1993; sity and patterns of bone loading, related to behav-
Hitchcock & Bleed, 1997). It is also the case that ioural variation in activity levels and activity patterns.
human use of hunting technology is highly variable The relationship between behaviour, bone loading and
from situation to situation (and hence highly adaptable modelling/remodelling is a complex one (see for
to circumstance), making it naive to talk about the example, Lanyon, 1990; Hylander & Johnson, 1997;
function of subsistence tools in any sort of absolute Daegling & Hylander, 1997; Demes et al., 1998) and
terms (see, for example, Hill & Hawkes, 1983; Bleed, we agree with Daegling (1993: 247) that ‘‘. . . the
1986; Greaves, 1997). However, among recent forag- inference of . . . stress patterns from purely morpho-
ers, hand-delivered spears tend to be thrust at prey logical criteria in the absence of experimental
from close quarters and, in the much rarer cases in corroboration is problematic’’. In this vein we
which they are thrown, tend to be delivered from close view behavioural models derived from analysis of
range (generally within 6 m: Churchill, 1993). Thus the skeletal morphology as hypotheses that must be tested
hand-delivered spear, whether thrust or thrown, is by experimental studies of living animals. The results
essentially a close range weapon, and herein lies the presented here represent an attempt to evaluate
limitations that this weapon imposes on prey selection the inferences about hunting behaviour that have
and choice of hunting tactic (Churchill, 1993). Spear been drawn for asymmetry and shape differences in
thrower-delivered spears, on the other hand, allow an Eurasian Neandertals and early modern human
effective hunting distance of 25–45 m (Churchill, 1993; humeri.
Cattelain, 1997), permitting greater flexibility in hunt- There is still considerable debate among bone biolo-
ing method and prey choice (although this weapon gists concerning the critical stimulus that triggers adap-
system has other limitations: see Churchill, 1993). Our tive modelling. Some argue that the critical stimulus is
central question, then, is not when spear throwing first the magnitude of the strain engendered in the bone
occurred in human evolution (as this behaviour un- during loading, not the number of loading cycles (e.g.,
doubtedly dates to the first spears!), but rather when Rubin & Lanyon, 1984; Whalen, Carter & Steel, 1988;
human hunters shifted from a focus on close range Beaupre, Orr & Carter, 1990a,b, see also Skerry &
predation with hand-held weapons to a focus on long Lanyon, 1995). It is certainly the case that even single
range hunting with projectile technology. The differing episodes of highly stressful activities have been shown
emphasis of these hunting modes on bimanual thrust- to stimulate bone remodelling (Chamay & Tschantz,
ing versus unimanual throwing provides hope that this 1972; see also Lanyon, Rubin & Baust, 1986). On the
transition will be reflected in the bony architecture of other hand many authors (e.g., Lanyon & Rubin, 1985;
human fossil upper limb remains. Martin & Burr, 1989; Rubin et al., 1994) do not believe
Spear Use in Pleistocene Homo 107
determine the contribution made by each limb to left limb in the trailing position. All subjects were
propelling the spear, to determine forces operating asked to thrust the shaft into a 1·25 cm thick stiff
along the spear shaft that are imparted to the limbs, gymnastics cushion mounted on a wooden frame. The
and to determine the limb positions of the subjects frame was secured against a wall so that it moved
during and after peak impact. These data are used to minimally. The 1·3 m2 cushion allowed a wide target
test the hypotheses, outlined above, derived from ob- area and thus provided the subjects considerable
servation of humeral diaphyseal cross-sectional mor- latitude in their posture and movement.
phological patterns in fossil humans. Essentially, Subjects were videotaped from a lateral view during
we endeavour here to provide middle range research spear thrusting with a single Panasonic CCD camera
(Binford, 1981; see also Churchill & Schmitt, 2001) (WV-D5100). Subjects were recorded from the right or
that provides an empirical link between structural left side depending on which limb was the trailing limb.
analysis of fossil long bones and inferences of prehis- This camera was electronically shuttered at 1/1000 s
toric behaviour. As stated above and in previous and images were recorded on a Panasonic AG-7350 at
publications (Churchill & Schmitt, 2001), we do not 60 images per s. Subjects wore bright reflective markers
assume that structural analysis of humeri will lead to at the wrist, elbow, and shoulder of the trailing limb
accurate behavioural inferences. Instead, we are testing (Figure 1). The spear was also marked with bright
hypotheses derived from previous structural analyses. white markers just behind the gauges. These markers
Because the results reported here deal only with exter- were used later for digitizing limb and spear position.
nal loads our data represent a first step in testing the Subjects were asked to perform three different thrust
validity of these hypotheses. types. A single brief strike followed by rapid with-
Two sets of four single element strain gauges were drawal (strike-withdrawal: SW), a series of repeated
positioned at two locations on the pole. The front set short thrusts and withdrawals (repeated strike-
was placed 240 mm from the distal end of the pole, the withdrawal: RSW), or a strike followed by a holding
back gauges were placed 750 mm from the distal end. period between one and three seconds (strike-hold:
Each set of four gauges were bonded around the shaft SH). Subjects were allowed to thrust the spear in any
equidistant from each other (such that if one were to way that they felt comfortable and were not restricted
look at the pole in section, the gauges would be concerning angle of attack and limb or body position.
positioned at 12 o’clock, 3 o’clock, 6 o’clock and 9 Signal output was in volts. The gauge output was
o’clock) using cyanoacrylate and covered with plastic calibrated after each trial using a known 1000 micros-
insulation (M-Coat D; Micromeasurements Corp.). train signal. These calibrations provided a value of 348
The gauges were connected in series and formed one (2·28) microstrain/volt. The calibration output from
arm of a Wheatstone bridge in a Vishay strain signal the back gauge were slightly lower (99·660·12%)
conditioner (Vishay #2110, #2120). The arrangement than the values for the front gauge, but these values
of the strain gauges allowed us to collect only measures were not significantly different. This reflects small
of axial (compressive) strains along the shaft. Bending differences in the amplifiers. The voltage signals from
moments produce strains of equal magnitude but each set of gauges were passed through an analogue-
opposite sign on opposite sides of the shaft. Thus by digital converter (Peak Performance Technologies,
arranging pairs of gauges opposite one another and Englewood, CO) and then read into Motus motion/
summing the strain signals, the tension and compres- kinetic analysis software (Peak Performance Tech-
sion aspects of bending cancelled each other out. To nologies, Englewood, CO). Data from the gauges were
confirm this we forcefully bent the pole by gripping on recorded prior to beginning the thrust and until the
either side of the gauges. Even with high bending forces strike(s) and the holding period(s) were complete and
the microstrain values were negligible. the spear was pulled off of the target.
Subjects were asked to grip the shaft in a fashion Since in linearly elastic materials there is a consistent
that was comfortable for them with one hand placed relationship between stress and strain (at least in the
between the front and back gauges and the other range of elastic deformation), we were able to estimate
behind the back gauges (Figure 1). Subjects were reaction forces (in Newtons) from observed micro-
allowed to hold the spear in any posture they chose and strain values. To do so, we placed the spear perpen-
were not instructed as to which limb should be in back. dicular to the platform of a Kistler force plate (Kistler
Seven subjects reported being right-handed and all Corporation) and applied loads with steady, strong,
seven used their right limb as the trailing limb. One rhythmic pushes. We collected 5294 paired measures of
subject was left-handed, and for three trials she used microstrain and compressive force using this approach.
her right limb as the trailing limb (the video camera The maximum voltage we achieved through this
was positioned for right-handed subjects, and initially method was 0·4 volts (139·2 microstrain), lower than
our left-handed subject conformed to this arrange- the voltage outputs commonly achieved during
ment). However, after a few trials our left-handed the trials (mean=0·56, max of 1·157 volts). Addition-
subject reported that it was uncomfortable to use her ally, this method did not entirely simulate the sharp
right hand as the trailing hand. Thus the camera was dynamic stabs our subjects used. Since estimating the
repositioned and for the remaining trials she used her forces associated with our highest observed microstrain
Spear Use in Pleistocene Homo 109
(a)
Red gauges (front), Raw analog data Green gauges (back), Raw analog data
Subject #3: Strike and hold
0.1
–0.1
Volts
–0.5
0.2 0.3 0.4 0.5 0.6 0.7 0.8
Time (sec)
(b)
Subject #3: Limb position at strike
Figure 2. (a) Raw analogue output from the front and back strain gauges during a strike and hold period. (b) A stick figure diagram produced
by digitizing reflective markers on the trailing limb at the point of impact. The angles calculated are indicated. In addition one moment arm
is illustrated. The moment arms for the shoulder, midshaft humerus, and wrist are not included in this figure.
values requires extrapolation beyond our empirical The ratio between the front and back gauges is a
regression, we also calculated the expected stress and convenient way to express which limb is providing
compressive force for a given strain based on Young’s most of the force of the thrust. If the force is applied
modulus of aluminum (70 Gpa) and the area of the from a single point behind both gauges then the output
metal within the cross section of the spear (121·7 mm). from both the front and back set of gauges will remain
The values for the equation from this approach yield equal. Tests of force applications to the pole in this
force values in Newtons that are close to, but slightly fashion confirmed this result. If the force is applied
higher than those of the empirically determined values only from behind the front gauges, the back gauges will
up to 139 ms (ratio of actual/predicted values= record no strain.
89·96%). Based on these procedures we are able During spear thrusting postures the force is applied
to provide some sense of the magnitude of load behind both sets of gauges; one hand behind the front
experienced during spear thrusting. gauges only and the other behind the front and
Typical kinetic and kinematic data are illustrated in back gauges (Figure 1). Thus the output from the front
Figure 2. The video data were recorded synchronously gauges will always be larger than the output from the
with the strain data using an Event and Video Control back gauges. If the force applied by both limbs (one
Unit (Peak Performance Technologies, Englewood, behind the front and one behind the back gauges) is
CO). The peak value for the front and back gauges equal then the output of the front set of gauges will be
were recorded for each strike and during each holding twice as large as that of the back gauges (back gauge/
period (Figure 2a). The ratio of back/front gauge front gauge ratio=0·5). In this situation, because of the
output was recorded at spear strike and during the alignment of the trailing limb with the spear and the
holding period. elevation of the forearm and arm from the spear
110 D. Schmitt et al.
Front gauges
Mean (..) 194·88 (73·1) 1660 (622·7) 37·9 (20·9) 3228 (178·0)
Range 57·8–402·6 492–3430 5·2–104·4 45–889
Back gauges
Mean (..) 123·54 (45·2) 1052 (385·4) 20·8 (10·6) 178 (90·4)
Range 44·2–248·1 377–2113 1·7–52·2 15–445
Back/front ratio
Mean (..) 0·647 (0·119) 0·585 (0·203)
Range 0·344–0·869 0·128–1·0
10
Motus. We were able to calculate estimates of the
magnitude of bending moments associated with the 8
spear reaction force at the wrist, elbow, shoulder, and 6
midshaft humerus. Moments were derived using the
limb and spear angles, the known lengths of the limb, 4
and by treating the spear as a force vector whose 2
orientation is given by the position of the spear and
0
whose magnitude is given by the compressive force (N)
5
5
5
.6
0
0
0
.6
.7
.8
.5
.7
.8
.5
.5
>0
>0
>0
>0
>0
>0
>0
<0
calculation is shown in Figure 2b. Because the highest Back gauge output/front gauge output
magnitude forces occurred at the instant of spear Figure 3. The distribution of gauge ratios (back gauge output/front
strike, the moments were calculated for this instant gauge output) for our 76 spear strikes. Note that only 7 of 76 were
only. less than 0·50 and 47 of 76 were equal to or greater than 0·60.
Spear Use in Pleistocene Homo 111
Table 4. Trailing limb and spear angles at spear strike Table 5. Moments of the spear force (Newton-meters) in the trailing
limb
Angle (degrees) Mean (..)
Mean (..) Range
Shoulder to X-axis 140·3 (7·1)
Elbow 101·6 (7·3) Wrist 137·0 (45·4) 103–223
Wrist 171·0 (8·6) Elbow 524·8 (152·8) 357–788
Spear to hand 114·4 (9·8) Mid-shaft humerus 732·5 (214·3) 500–1118
Shoulder 888·4 (309·6) 574–1374
It is likely that hunters would often encounter resist- about the shoulder (Table 5) are a little more than half
ance from bone during penetration. We anticipate that those experienced in torsion during throwing—
impact forces when a spear encounters bone or other approximately 890 Nm during thrusting versus
substantial resistance are similar to those generated by 1665 Nm of torsional force during throwing (Gainor
our subjects in the lab, and that our experimental et al., 1980). Nonetheless, the loads during thrusting
approach provides an initial understanding of the ways are likely more than adequate to stimulate osteogenic
that forces are imparted to the two limbs during activity to a greater degree in the dominant (trailing)
bimanual spear use. limb versus the nondominant (leading) limb.
During the strike period the subject must resist The most salient point about the moments during
moments tending to retract the shoulder, extend the spear thrusting in the dominant limb is the orientation
elbow, and ulnarly deviate the wrist of the trailing of the bending moments. The moments along the
limb. Subjects generally must recruit muscles to resist forearm occur in the mediolateral plane because the
turning moments at these joints. The insertion sites of volar surface of the forearm faces medially. However,
muscles that resist the reaction forces generated by the bending moments along the humerus appear to
spear thrusting, notably the humeral flexor M. pecto- occur primarily in the anteroposterior plane, consistent
ralis major and the forearm flexors Mm. biceps brachii with the pattern of bone cross-sectional anatomy seen
and brachialis, are hypertrophied in Neandertals in Eurasian Neandertals and Early Upper Paleolithic
(Trinkaus, 1983a,b) and in some early modern Euro- modern humans.
peans (Churchill & Formicola, 1997; Churchill & It should also be noted that the guiding (leading)
Smith, 2000). Subjects may also rely on osteoligamen- limbs may also experience high moments. This limb
tous structures to help resist turning moments at the projects somewhat laterally from the long axis of the
shoulder, elbow and wrist. Along these lines it is spear and as a result the humerus of this limb may have
interesting that at the moment of spear strike (and experienced considerable torsion. On the other hand,
hence peak reaction forces) our subjects had partially because the limb is held strongly flexed, the action of
flexed elbows (average angle 101·6: Table 4). As biceps and triceps along the shaft may lead to relatively
Trinkaus & Churchill (1988: 20) noted in reference higher anteroposterior loads. This may explain the
to the morphology of the ulnar trochlear notch in similar shape of the right and left humeri (Table 1)
Neandertals: whereas the strength asymmetry is arguably
Most of the Neandertal ulnar trochlear notches are more explained by the asymmetry in the magnitude of loads
anteriorly oriented than those of more recent humans applied.
(Trinkaus, 1983a). This proximal ulnar articular orien- Taken together, the asymmetry of load and the
tation of the Neandertals would provide greater resistance orientation of the bending moments support in-
to joint reaction forces in partial flexion of the elbow (by
ferences drawn from skeletal anatomy that Eurasian
maximizing the effective joint surface area perpendicular
to the joint reaction force and the ability of the joint to Neandertals and Early Upper Paleolithic hunters relied
resist proximodistal shear stress), whereas the more proxi- heavily on thrusting spears and close-range hunting.
moanterior orientation of recent human ulnar trochlear The humeri of males from Mousterian and Early
notches would more effectively resist such forces in exten- Upper Paleolithic contexts exhibit a right-dominant
sion (this in no way implies any difference in ranges or pattern of strength asymmetry, and appear to have
patterns of elbow movement, only a frequency difference been well prepared to withstand bending moments in
in the habitual position of peak loading). the parasagittal (anteroposterior) plane. Coupled with
Another way that a subject may control bending the lack of direct evidence for projectile weaponry
moments on long bones and compressive and shear during this time, we feel the most parsimonious con-
forces at joints is by positioning the limb such that the clusion is that thrusting spear use was one of the
bones are more in line with the force vector and joint principle sources, if not the principle source, of osteo-
surfaces are perpendicular to it. We found our subjects genic stimuli in Neandertal and Early Upper Paleo-
to be fairly conservative in the postures chosen to lithic modern human male humeri. Males of the Late
strike the target (both between trials of the same Upper Paleolithic tend to be characterized by humeri
subject and between subjects). This suggests that the that are more equally resistant to bending moments in
best posture for generating thrusting force and resist- multiple directions (and torsion, as generated during
ing reaction forces may fall within a fairly narrow throwing) as well as right-dominated strength asym-
range of posture. Regardless, subjects still engendered metry. In conjunction with artifactual evidence of
large bending moments in the trailing limb. The high projectile weaponry in the Late Upper Paleolithic, we
loads along the long axis of the spear were coupled suggest that torsional moments generated during hunt-
with long moment arms, some of which were greater ing with long-range projectile weapons was one of the
than 75% of the length of the fully extended limb principle sources of osteogenic stimuli in males of this
segment. This resulted in bending moments as high or period. Thus we feel, based on current evidence, that
higher than those seen in the lower limb during walk- long-range projectile weaponry did not become a regu-
ing and distance running (Andriacchi, Natrajan & lar part of human predatory methods until the early
Hurwitz, 1997). The bending moments we observed part of the Late Upper Paleolithic (Solutrean), roughly
Spear Use in Pleistocene Homo 113
coincident with the first appearance of spear-throwers Exploitation in the Later Palaeolithic and Mesolithic of Europe.
in the archeological record. American Anthropological Association Archaeological Papers,
vol. 4, pp. 11–24.
Churchill, S. E. & Formicola, V. (1997). A case of marked bilateral
asymmetry in the upper limbs of an Upper Paleolithic male from
Acknowledgements Barma Grande (Liguria), Italy. International Journal of Osteo-
archaeology 7, 18–38.
We are very grateful for all the help we had with this Churchill, S. E. & Smith, F. H. (2000). A modern human humerus
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important comments and advice throughout. We thank study of behavioural evolution in the genus Homo. In (C.
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