Humans, tools and handedness

James Steele* and Natalie Uomini

#


* AHRC Centre for the Evolution of Cultural Diversity, Institute of Archaeology, University College
London, 31-34 Gordon Square, London WC1H 0PY, UK. Email: j.steele@ucl.ac.uk

#
Department of Archaeology, University of Southampton, Highfield, Southampton SO17 1BJ. Email:
n.t.uomini@soton.ac.uk

Published in: Stone Knapping: The Necessary Conditions for a Uniquely Hominin Behaviour
(edited by Valentine Roux and Blandine Bril), pp. 217-239. Cambridge: McDonald Institute
for Archaeological Research, 2005.
217
Humans, Tools and Handedness
ChaIer 15
Humans, Tnn!s and Handcdncss
cal inleresls. We shall lhen discuss some skelelal and
malerial-cullural markers of hand reference lhal may
enable us lo lrack lhe evolulion of human handedness
emirically.
MeasurenenI of handedness
Handedness is measured in a variely of vays, vhich
are eilher reference based or erformance based
(McManus 1996). Skill and reference lend lo be
highly correlaled, allhough lhere are excelions. Skill
is usually measured by comaring lhe lvo hands
in raid aimed movemenl lasks such as lhe Annell
egboard lask (Annell 197O), vhich measures hov
fasl each hand can move len egs in a board from one
rov of holes lo anolher. Handedness is assessed by
calculaling lhe relalive seed advanlage of lhe more
skilled hand, and is lherefore lrealed as a conlinuous
variable. Ireference is usually measured by queslion-
naires vhich ask aboul lhe referred hand for each of
a series of lasks, and in vhich lhe resondenl indicales
lhe slrenglh of lhe reference on an ordinal scale (e.g.
Alvays Righl, Usually Righl, Lilher, Usually Lefl,
Alvays Lefl). Resonses are usually summed inlo a
Laleralily Index by allocaling values (e.g. +2, +1, O,
1, 2) lo each osilion on such a scale. In younger
children and non-human rimales, for vhom a ques-
lionnaire is inaroriale, reference may inslead be
direclly observed in a series of simle lasks.
olh measures give comarable resulls vhen
used lo delermine broad allerns of lefl- and righl-
}ames Sleele & Nalalie Uomini
Prccisicn icc| usc iqpicc||q intc|tcs prcjcrcniic| usc cj c !cmincni ncn! in numcns cn!
scmc cincr cnimc| spccics, cn! in numcns inc rigni ncn! icn!s ic |c inc prcjcrrc! ncn!. Wc
cui|inc ccntcniicnc| criicric jcr rcccgnizing ncn!c!ncss in |iting su|jccis, cn! summcrizc
scmc rcccni gcnciic mc!c|s cj inc sic|i|iiq cj numcn ncn!c!ncss cs c pc|qmcrpnism. Wc
incn summcrizc skc|cic| cn! (in grccicr !cici|) mcicric|-cu|iurc cti!cncc jcr ncn! prcjcrcncc
in inc jcssi| cn! crcnccc|cgicc| rcccr!s. Sucn c|scrtciicns suggcsi inci rigni ncn!c!ncss
ncs |ccn prc!cmincni ctcn in ccr|q spccics cj cur cun gcnus, c|incugn inc jcssi| scmp|c
is cxccc!ing|q smc||.
Thc rn!cs nI thc hands In tnn!-usIng tasks
Skilled lool maniulalion usually falls inlo lhe
calegory of asymmelric or differenlialed bimanual
aclivilies (Guiard 1987, 487). Wilh remarkable con-
sislency, individuals divide lhe vork belveen lheir
lvo hands in a rediclable and regular fashion. More
secihcally, one hand lends lo be referred as lhe one
lhal execules an aclion on lhe ob|ecl, vhile lhe olher
hand slabilizes lhe ob|ecl. Mosl remarkably, in aboul
eighl or nine oul of len individuals il is lhe righl hand
lhal is selecled lo lay lhe leading role (making il lhe
dominanl hand). This role lyically involves hner
movemenls, in lerms of bolh salial and lemoral res-
olulion (Guiard 1987, 497). The allern is exemlihed
by slone knaing, vhere lhe dominanl hand vields
a hammerslone lo slrike a core lhal is suorled by
lhe non-dominanl hand.
This allern of oulalion-level hand reference
seems lo dislinguish us from olher living rimales,
among vhose oulalions and secies il is hard lo
discern any such bias. ecause our ovn righl-handed
bias seems lo be relaled, via lhe linking mechanism of
cerebral dominance, lo anolher unique human fealure
(language rocessing: e.g. Hcaen & de A|uriaguerra
1964, radshav & Rogers 1993), an enormous volume
of research has been dedicaled lo ils underslanding.
In lhis chaler ve shall inlroduce some asecls of
lhe research lileralure in sychology and behaviour
genelics, lo rovide conlexl for our ovn archaeologi-
218
Claplor 15
hand use al lhe oulalion level. The reference
measure has been more videly used, erhas because
il is simler lo adminisler. Inevilably some queslion-
naire ilems are sub|ecl lo cullurally-learned biases,
a faclor lhal obviously comlicales inlerrelalion of
lhe mosl simle reference measure used in lilerale
socielies (lhe vriling hand). Hovever, lhere is some
vell-designed research using bolh skill and refer-
ence measures, vhich indicales lhal queslionnaire
resonses can be analyzed and a faclor idenlihed
relaling lo recise molor conlrol vhich is imervious
lo cullural inuences (Connolly & isho 1992).
ne grou of researchers has allemled lo collecl
dala in an elhological manner, resembling lhe vay
handedness dala is collecled for non-human rimales
such as chimanzees. They found lhal human manual
reference in non-lool-using lasks is less righl biased
lhan imlied by lhe queslionnaire measures (vhich
are heavily biased lovards ob|ecl maniulalion lasks).
Marchanl ci c|. (1995) labulaled hand use observed in
elhnograhic videos from grous of lhree lradilional
cullures (G1vi, Yanomamo, and Himba) using lhe
kind of lask classihcalion lhal vould be used in an
ecologically-valid rimale sludy. Lxamles of behav-
ioural calegories of limb movemenls included reach-
ing for ob|ecls, scralching oneself, ealing, and using
lools. Their resulls evidenced a barely-discernible
righl hand reference for all lasks al lhe oulalion
level, allhough a slronger righl-hand bias vas found
for lool use only (arlicularly vhere il involved a
recision gri, vhich agrees vilh Guiard 1987).
AsnneIrles ln hand sklII and notenenI conIroI
In exlaining lhe funclional neurology of human
handedness, mosl researchers lake lhe righl-hander as
lheir rololye. Hand skill is oflen measured by aimed
movemenl lasks (such as lhe Annell egboard lask,
described above), and righl-handedness is generally
exlained by reference lo a lefl hemishere advanlage
for hne lemoral resolulion of sensory inul and molor
oulul (Carson 1993). An advanlage for lhe dominanl
hand is usually seen nol in simle ballislic movemenls,
bul in movemenls of grealer difhcully (in lhe Iills'
Lav sense, Iills 1954, also in lerms of hner salial and
lemoral resolulion, Guiard 1987). Ilovers (1975) hy-
olhesized lhal in movemenls of grealer difhcully as
measured using Iills' Index, and vhich imly a 'correc-
live mode of conlrol', lhe dominanl hand vould have
an advanlage because of an underlying advanlage in
lhe rale of informalion lransmission. Carson (1993, 481)
discusses lvo exlanalions for lhis advanlage. ne is
lhe 'feedback rocessing' model, vhich rooses lhal
lhe lefl hemishere is more efhcienl in error correclion
using sensory feedback. The olher is lhe 'oulul vari-
abilily' model, vhich rooses lhal lhe lefl hemishere
ermils more recise conlrol of nel forces and force du-
ralions. There is slill considerable uncerlainly regarding
vhich of lhese models is more valid.
There is also considerable debale regarding lhe
level of organizalion al vhich neurohysiological
asymmelry is found. A volunlary bimanual movemenl
can be analyzed in lerms of lhree levels of organizalion
(Ielers 1995, 2O1). These are Level 1 (lhe level al vhich
lhe goal is formulaled), Level 2 (lhe level al vhich
'lhe recisely limed commands for lhe inilialion and
lerminalion of lhe movemenl lra|eclories of lhe lvo
hands are issued': Ielers 1995, 2O2), and Level 3 (lhe
level 'vhich governs lhe hnal oulov of conlrol for
lhe arlicular hand lhal allovs lhe hand lo erform
lhe movemenl as required': Ielers 1995, 2O3). Ielers
(1995) favours an asymmelry in allenlional rocesses
vhich inuences hand skill al Level 1, vhile Sainburg
(2OO2) suggesls lhal lhe causal agenl is an asymmelry
in lhe conlrol of limb segmenl inerlial dynamics vhich
occurs dovnslream from lhe lra|eclory lanning level.
Il is beyond lhe scoe of lhis chaler lo do more lhan
nole lhese debales, and oinl oul lheir relevance lo
underslanding lhe kinds of skilled movemenl conlrol
involved in lhe knaing geslure (e.g. Roux ci c|. 1995,
Roux 2OOO).
Exp!anatInns nI thc prcva!cncc nI !cIt- and rIght-
handcdncss
The observalion lhal olher living rimales do nol
shov as slrong a righl-handed bias as humans al lhe
oulalion level (MacNeilage ci c|. 1987, Marchanl
& McGrev 1991, Sugiyama ci c|. 1993) has led some
evolulionary sychologisls lo con|eclure lhal lhe
inilial ralio of righl and lefl-handedness vas 5O:5O
in early hominins. If lhis is correcl, lhen lhe resenl
ralio of aboul 9O:1O in humans can only have arisen
subsequenlly lhrough nalural seleclion. This imlies
lhal righl-handed individuals had a reroduclive
advanlage, namely lhal lhe genes associaled vilh
righl-hand dominance vere osilively selecled for and
vere able lo sread via Mendelian inherilance lhrough
our secies. Irom lhis argumenl il also follovs lhal
some exlanalion musl be given for lhe ersislence of
a small roorlion of lefl-handers, desile lhis selec-
lion lovards righl-handedness.
There are several comeling exlanalions for lhe
resenl-day incidence of righl- and lefl-handedness,
and lhree of lhese vill be delailed belov. Some os-
lulale lhal human righl-handedness is lhe norm, and
lhal lefl-handedness is alhological, hovever, lhere is
lillle emirical suorl for such an exlreme osilion.
lhers have roosed lhal vhile lhere are disadvan-
219
Humans, Tools and Handedness
lages lo lefl-handedness, comensaling advanlages
may accrue vhich are frequency-deendenl. Mosl
commonly, hovever, lhe argumenl is made lhal hand-
edness is under arlial genelic conlrol vilh signihcanl
environmenlal modihcalion during develomenl. In
such models, il is argued lhal lhe advanlage lies vilh
lhose vho are moderalely righl-handed, bul lhal lhe
inleraclion of genelic and environmenlal inuences
roduces grealer henolyic varialion, ranging from
lefl-handedness lo exlreme righl-handedness. All such
exlanalions have lo address lhe emirical hndings of
an aarenlly slable underlying revalence of aboul
1O15 er cenl lefl-handedness, and of a male excess
(aboul hve males are lefl-handed for every four fe-
males: McManus 1996).
The mosl famous of lhe exlanalions vhich
roose lhal lefl-handedness is alhological is lhe
Geschvind-ehan-Galaburda hyolhesis (Geschvind
& ehan 1982, 1984, Geschvind & Galaburda 1985a,b).
This rooses lhal individuals, by defaull, develo lo
be righl-handed unless lhere is some lesloslerone-in-
duced develomenlal delay in lhe grovlh of lhe felal
lefl hemishere, causing nol |usl lefl-handedness bul
also alyical language laleralizalion (and olher less
inluilive disorders, such as a high rale of auloimmune
disease). This hyolhesis has been exhauslively exam-
ined in clinical sludies, and lhe verdicl musl nov be
lhal il is nol suorled (e.g. ryden ci c|. 1994). lher
exlanalions exisl vhich relale lefl-handedness lo
develomenlal neurological disorders, arlicularly
in lhe conlexl of felal grovlh relardalion and of re-
malure birlh (akan 1971, akan ci c|.1973, Salz 1972).
While such lrauma, hovever, may accounl for a small
fraclion of lefl-handers vhose reference is genuinely
secondary lo felal brain insull, il does nol seem lo
exlain lefl-handedness in more lhan aboul one in
lvenly cases (isho 1984).
A second class of exlanalion inlerrels lhe er-
sislence of lefl-handedness in lov frequencies as due lo
some cognilive or olher advanlage, vhich counleracls
any develomenlal disadvanlages. This only vorks
if lhe frequency of lefl-handedness slays belov some
crilical level. The evidence for an associalion belveen
develomenlal delay and elevaled frequencies of non-
righl-handedness is quile slrong, vilh some sludies
indicaling grealer risk for shorl slalure, reduced body
mass, and delayed onsel of uberly (e.g. Coren &
Halern 1991, Mulligan ci c|. 2OO1, bul see also Lalon
ci c|. 1996). Claims for some secihc comelilive ad-
vanlage associaled vilh lefl-handedness are based on
aarenlly elevaled frequencies of lefl-handedness in
cerlain aclivilies and rofessions (Ielerson & Lansky
1977, bul see Wood & Agglelon 1991, Meberl & Michel
198O, Goleslam 199O). More recenlly, an argumenl for
a frequency-deendenl advanlage for lefl-handed
hghling has been roosed, based on observed el-
evaled incidences of lefl-handedness in inleraclive
and combal sorls (Raymond ci c|. 1996). In arlicular,
because lefl-handers are in lhe minorily, lhey are more
successful vhen lhey hghl againsl righl-handers. Such
roosals enlail secihc rediclions aboul lhe inlerac-
lion belveen cullurally-variable seleclion coefhcienls
for lefl-handedness, and lhe dislribulion of variance in
reroduclive success: higher hlness vould be execled
for lefl-handers vho live in very violenl socielies. Hov-
ever, delailed genelic models lo suorl such roosals
have nol yel been arliculaled, and unlil lhey have been,
ve should be vary of confusing correlalion vilh causal
exlanalion.
The lhird class of exlanalory models invokes
lhe genelic lheory of a balanced olymorhism vilh
helerozygole advanlage. The single-locus lheories for
laleralily roose lhal lhere is a gene, made u of lvo
alleles (eilher one can be recessive or dominanl), for
lefl-hemishere cerebral dominance, vhich causes
slrong righl-handedness as vell as language lalerali-
zalion. Annell (1985, 2OO2) calls lhis lhe Righl Shifl
allele (R+), and lhe allernalive is simly an inaclive
allele (R, vhich ve vill refer lo here as O). Since each
erson inherils one allele from each arenl, lhere are
lhree ossible genelic combinalions (genolyes): 2
R alleles (homozygous), 2 O alleles (homozygous),
or 1 R allele and one O allele (helerozygous). In lhe
absence of lhe R gene, individuals may develo righl-
or lefl-hemishere dominance vilh equal likelihood
as a resull of chance environmenlal faclors during
develomenl (Annell 1985, 2OO2, McManus 1985,
cf. Laland ci c|. 1995). The reason vhy ve do nol
all have lhis gene is because lhere is an advanlage
for being helerozygous (RO) al lhis single locus. Il is
beller lo have one allele causing lhe 'righl shifl', and
anolher allele vhich gives no such bias, because lheir
inleraclion vill lend lo roduce lhe olimal oulcome
moderale lefl-hemishere dominance, and lhus,
moderale righl-handedness. Some individuals (OO
homozygoles) vill have a comlele absence of lhe
righl-shifl gene (vhich, cullural biases exceled, vill
lend lo roduce lefl-handedness in aboul half of lhe
cases), and olher individuals (RR homozygoles) vill
have a double dose of lhe righl-shifl gene (vhich vill
bias lovards exlreme righl-handedness).
The idea of a helerozygole advanlage is nol nev,
lhe classic examle being lhe malaria hyolhesis for
sickle-cell anaemia (Haldane 1948, ciled in Durham
1991, 123). Among lhe oulalions of lroical Wesl Af-
rica lhere are lhree classes of haemoglobin genolyes:
AA, AS, and SS. The A allele is lhe normal condilion
for haemoglobin. The recessive S allele is a mulalion
22O
Claplor 15
of lhe haemoglobin molecule vhich causes sickle-cell
anaemia, bul also confers resislance lo malaria. Ieole
vilh lhe AA genolye have a normal (severe) reaclion
lo malaria, eole vilh lhe SS genolye have severe
sickle-cell anaemia, bul eole vilh lhe AS genolye
only shov very veak sickle-cell symloms and very
lov rales of malarial infeclion and morlalily. The ad-
vanlage of lhe AS genolye lies in lhe combinalion of
normal and sickling haemoglobin. AS haemoglobin,
only vhen infecled vilh malaria, begins a sickling roc-
ess vhich leads lo lhe dealh of lhe malarial arasiles.
Normal AA haemoglobin does nol have lhis caabilily,
vhereas lhe anli-malarial abilily of SS haemoglobin is
overshadoved by lhe high morlalily caused by sickle-
cell disease (Durham 1991, 1O6 ff. & 481).
These laller genelic models do quile vell in ac-
counling for lhe allerns ve observe for herilabilily
of handedness, and for lhe allerns observed in lvin-
ning. To dale, hovever, lhe evidence for a quanlihable
helerozygole advanlage associaled vilh moderale
righl-handedness has been equivocal. Several al-
lemls lo idenlify lhis advanlage have invesligaled
ils ossible behavioural origin (i.e. a cognilive advan-
lage: Annell & Manning 1989, a link lo schizohrenia:
Crov ci c|. 1998, bul see Nellle 2OO3), allhough lhere
have been fev if any sludies of lhe measurable direcl
effecls on reroduclive hlness. Recenl vork by Yeo
& Gangeslad (1993, Yeo ci c|. 1993) does advance lhis
held somevhal, allhough slill nol measuring direcl
hlness consequences of helerozygosily al lhe cerebral
dominance locus. They have found lhal comared
vilh moderalely righl-handed individuals, bolh
lefl-handers and exlreme righl-handers have higher
incidences of lhe minor hysical anomalies lhal are
associaled vilh develomenlal inslabilily (vhich is, in
lurn, associaled vilh generalized homozygosily).
Skc!cta! cnrrc!atcs nI handcdncss
To summarize our discussion so far, il seems lhal lhere
is an advanlage for lhe dominanl hand in lool use lhal
relales lo an underlying efhciency in informalion-lrans-
fer rale in lhe conlralaleral cerebral hemishere. This
advanlage is seen in lhe grealer skill of lhe dominanl
hand vhen execuling volunlary movemenl lasks vilh
high levels of difhcully in lhe Iills's Lav sense, or lasks
vilh very hne salial and lemoral resolulion in lhe
Guiard sense, and in vhich a correclive mode of conlrol
is indicaled. Il seems likely lhal lhe neurological basis
of lhis asymmelry in skill is only veakly genelically
delermined, vilh considerable scoe for environmenlal
inuence during develomenl. Individuals vary bolh
in lheir hand reference and in lhe degree lo vhich one
hand is more skilled lhan lhe olher.
The evolulionary origins of lhe mosl commonly
observed allern, namely a lefl-hemishere seciali-
zalion for lhe execulive role and consequenl righl-
handedness, are of course mallers of inlense scienlihc
inleresl and debale. Il is beyond lhe scoe of lhis
aer lo reviev such debales furlher, since lhal vould
require us lo digress inlo lhe evolulionary analomy
of language. In lhis seclion, lherefore, ve shall sum-
marize some forms of skelelal evidence enabling us lo
lrack lhe evolulion of handedness in lool use as seen
in lhe fossil record (see also Sleele 2OOO, from vhich
lhis lhree-age summary is abslracled).
Many sludies in recenl years have demonslraled
lhe range of adalive resonses of lhe skelelon lo al-
lerns of mechanical loading in .i.c (e.g. Carler 1987)
These resonses can include increases in bone slrenglh
due lo increased bone densily and1or cross-seclional
area, increases in mechanical efhciency by shae
change, and resislance lo avulsion by increasing lhe
surface area of lhe siles of allachmenl of muscles and
ligamenls on a bone's surface. Lvidence suggesls lhal
in any arlicular case, lhe effecl of muscle slrenglh and
mechanical loading on bone-mineral formalion is lo-
calised lo lhe secihc sile of musclebone inleraclion.
ecause a consislenl hand reference leads lo laleral
asymmelry in lhe mechanical loading exerienced by
lhe lvo hands, arms and shoulders during life, ve can
diagnose lhe handedness of a deceased individual by
sludying righllefl differences in lhe lifelime skelelal
resonse lo loading slrains.
In humans, suorling evidence is found from
a number of sludies lhal have quanlihed skill and
slrenglh differences belveen lhe dominanl and lhe
non-dominanl hand and arm. Il is lausible lhal some
skill differences are develomenlally canalized, bul
lhal skill and slrenglh differences are subsequenlly
amlihed by habilual allerns of use. Annell has
found lhal in lhe egboard lask, lhe dominanl hand
is caable of erforming vilh an average seed ad-
vanlage over lhe olher hand of 4.2 er cenl in females
and 3.4 er cenl in males (Annell 1998). lher similar
lasks roduce larger skill asymmelries belveen lhe
lvo hands (1O12 er cenl: Taley & ryden 1985).
Il has also reealedly been observed lhal in righl-
handed adulls of bolh sexes, normal gri slrenglh
in lhe dominanl hand lends lo be aboul 1O er cenl
grealer lhan lhe gri slrenglh of lhe non-dominanl
hand (Thorngren & Werner 1979, Ielersen ci c|. 1989,
Crosby ci c|. 1994, Chau ci c|. 1998). A similar allern
of relalive inch slrenglh belveen lhe dominanl and
non-dominanl hand has also been reorled (vilh lhe
dominanl hand aboul 1O er cenl slronger lhan lhe
non-dominanl hand: rorson ci c|. 1989, imson ci c|.
1997, Chau ci c|. 1998). Iurlher conlrasls relaling lo
221
Humans, Tools and Handedness
hand reference have been described for vrisl exlen-
sion, vilh lhe dominanl side having on average aboul
1O er cenl grealer vrisl exlension slrenglh (Richards
ci c|. 1993). Ierlexingly, hovever, lefl-handed sub|ecls
are equally likely lo have a slronger gri in eilher hand
(Crosby ci c|. 1994, Ielersen ci c|. 1989), erhas lhis
reecls lhe need for lefl-handers lo adal lheir hand-
use allern lo lhe conslrainls of a righl-handed vorld.
An effecl of vork allern has been reorled (}osly ci
c|. 1997): heavy manual vorkers have lhe slrongesl
gri and lhe leasl slrenglh difference belveen lhe
lvo hands, vhile ofhce vorkers have lhe veakesl
gri and lhe grealesl slrenglh difference belveen lhe
lvo hands. Lighl manual vorkers vere found lo be
inlermediale belveen lhese lvo grous.
The bones of Ihe hand, arn and shouIder glrdIe
A reealed clinical observalion is lhal lhe bones of
lhe righl hand lend lo be larger lhan lhose of lhe lefl
hand, as does lhe volume of lhe hand ilself (Iurves
ci c|. 1994). This is seen in radiograhs (McLeod &
Couland 1992), allhough in lhe earliesl sludies of
side differences in lhe second melacarals by Garn ci
c|. (1976) and by Ilalo ci c|. (198O) no correlalion vas
found vilh handedness (as measured eilher by hand
reference or by gri slrenglh). A correlalion of rela-
live hand size vilh handedness has been reorled for
righl-handers, allhough nol for lefl-handers (Iurves ci
c|. 1994). Mosl recenlly, Roy ci c|. (1994) have reorled
hnding bilaleral asymmelry of bone area in lhe sec-
ond melacaral correlaled vilh hand dominance, in
vhich handedness vas assessed by lhe sub|ecl's ovn
ersonal imressions of vhich handedness grou
lhey belonged lo.
In righl-handed adolescenls and adulls, muscle
mass lends lo be grealer in lhe arm on lhe dominanl
side (Chhibber & Singh 1972, Schell ci c|. 1985, Neu-
mann 1992, Taaffe ci c|. 1994). Il is also vell eslablished
lhal lhe righl humerus and radius lend lo be slighlly
longer and heavier lhan lheir lefl counlerarls (La-
limer & Lovrance 1965, Ruff & }ones 1981). There
have been a number of recenl radiograhic sludies of
rofessional racquel sorls layers and olher alhleles,
vho may begin lheir lraining early in childhood, and
vhose dominanl arms lend lo exerience unusu-
ally large mechanical loads during lhe laying years
(uskirk ci c|. 1956, }ones ci c|. 1977, Haaasalo ci c|.
1994, Tsu|i ci c|. 1995, Kannus ci c|. 1996). These slud-
ies have concenlraled on differences belveen lhe long
bones of lhe lvo forelimbs in bone-mineral conlenl,
bone mineral densily and cross-seclional corlical area.
They reealedly observe grealer bone-mineral densily
and conlenl in lhe long bones (humerus, radius, ulna)
on lhe dominanl side (a allern also found, bul less
markedly, in normal conlrol samles). Such modern
radiograhic sludies converge on lhe hnding lhal
aclivily slresses roduce adalive resonses in lhe
bones of lhe dominanl forelimb, effecls lhal oughl lo
be discernible as measurable asymmelries in aired
skelelal elemenls in individuals from archaeological
oulalions.
Asymmelrical loading allerns are also found in
eole vilhoul inlensive sorls lraining. Ingelmark,
in an early and ioneering radiograhic sludy (1946),
found lhal grealer forelimb lenglh (as measured by
lhe sum of lhe lenglhs of lhe humerus and radius)
vas correlaled vilh lhe side of lhe referred hand in
children. In lhis sludy he classihed as lefl-handers all
children vho reorled lhe use of lheir lefl hands in
al leasl lvo of seven everyday lasks. Consislenl vilh
modern behavioural dala on handedness, females are
also more likely lhan males lo have longer long bones
in lhe righl forelimb. Tvo recenl sludies of lennis lay-
ers and normal conlrols go some vay lo relicaling,
among adulls, Ingelmark's hnding (Krahl ci c|. 1994,
Haaasalo ci c|. 1996).
Reichel ci c|. (199O), in a radiograhic sludy
of normal adulls, also found a correlalion belveen
handedness and lhe side of grealer bone-mineral
densily and bone vidlh in lhe radius in ils midshafl
and dislal segmenls. The ulna aears lo be lhe bone
vilh leasl bilaleral asymmelry of lhe lhree long bones
in lhe arm. In rofessional racquel sorls layers,
lhe effecl of rolonged unilaleral loading on increase
in bone-mineral conlenl and bone mineral densily
in lhese bones is slighlesl al siles in lhe ulnar shafl
and lhe dislal ulna (Haaasalo ci c|. 1994, Kannus
ci c|. 1996). Iresumably lhis reecls ils lesser role
in dislribuling mechanical load in racquel sorls.
Kennedy (1983), hovever, has observed referenlial
develomenl of lhe ulnar suinalor cresl in lhe righl
arms of males in some archaeological oulalions of
modern humans, aarenlly reecling lhe slresses
involved in overarm lhroving (as, for examle, of a
hunling sear).
Lffecls of aclivily on lhe bones of lhe shoulder
girdle (scaula, clavicle) have been less frequenlly
sludied lhan lhey have on lhe long bones of lhe fore-
limb, bul all lhese bones oflen shov clear asymmelries
relaled lo handedness. Il is usual lo hnd a grealer
range of molion in lhe gleno-humeral |oinl (vhere
lhe humerus arliculales vilh lhe shoulder blade) on
lhe side of lhe referred hand (onci ci c|. 1986). In
lhe clavicles, lhe righl bone lends lo be bolh shorler
and more robusl lhan lhe lefl. Mays ci c|. (1999), in a
sludy of lhe clavicles from lhe redominanlly medi-
aeval oulalion of Wharram Iercy, have found lhis
same allern and have also found a lendency for lhe
222
Claplor 15
areas of allachmenl of lhe cosloclavicular ligamenl
(lhe sile of a fealure knovn as lhe rhomboid fossa)
and of lhe lraezoid ligamenl lo be more develoed
on lhe righl side. These aulhors suorl lhe hyolh-
esis lhal loading of lhe dominanl limb exerls grealer
axially comressive forces on lhe isilaleral clavicle,
leading bolh lo shae changes (grealer robuslicily)
and lo grealer develomenl of lhe allachmenl siles of
lhose ligamenls vhich slabilize lhe clavicle vilhin lhe
shoulder girdle during axial comression.
These osl-cranial skelelal markers of asym-
melrical develomenl in lhe shoulder, arm and hand
aear lo rovide us vilh a very exlensive lool kil for
diagnosing handedness in asl oulalions. Hovever,
a number of olher rocesses may affecl lhe develo-
menl of asymmelry in aired skelelal elemenls in lhe
forelimb and shoulder girdle, and lhese should also
be laken inlo accounl in any such analysis (cf. Sleele
2OOO, 21314). These olher rocesses include bolh
uclualing asymmelry, and direclional asymmelries
favouring grovlh in one member of a air of bones
vhen lhese are due lo innale develomenlal biases
and nol lo mechanical loading hislory. Ialhological
develomenl of elemenls of one side of lhe body is a
lhird olenlially comlicaling variable.
SkeIeIaI narkers of handedness ln hunan etoIuIlon
and rehlsIor
If our inferences aboul lhe relalionshi belveen hand-
edness, lool use and lhe adalalion of bone lo load-
ing are correcl, lhen ve vould execl asymmelrical
skelelal develomenl lo occur only in rimale secies
vhich are exlremely lool-deendenl (i.e. humans,
and lheir lool-deendenl hominin anceslors). Schullz
(1937) recorded asymmelries of lhe lenglhs of arm
bones (humerus and radius) in a large samle of ae
skelelons (including 13O gorillas, 82 chimanzees, 8
orangulans, and 21 gibbons). In marked conlrasl vilh
lhe 722 human skelelons in his samle, he found no
lendency for lhe righl arm lo be dominanl in aes as
assessed by lhis measure. He also found lhal lhe mean
degree of asymmelry (unsigned) in aes vas aboul
half lhal found in lhe arm bones of humans. These
hndings concur vilh lhe observalion, menlioned
above, lhal aes do nol exhibil eilher lhe oulalion-
level righl-handedness seen in humans, or lhe degree
of loading of lhe individually-dominanl side vhich is
seen in lhe human bones.
If ve examine skelelons of relalively recenl
oulalions from lhe hislorical eriod, lhe allerns
suggesl lhal frequencies of righl- and lefl-handedness
have been relalively slable across lhe cenluries. Sleele
& Mays's (1995) sludy of asymmelry in lhe summed
lenglhs of lhe humerus and radius in lhe medieval
Wharram Iercy cemelery oulalion found a allern
in adulls very similar lo lhal reorled by Annell for
lhe dislribulion of manual erformance asymmelries
in lhe modern rilish oulalion, vilh 81 er cenl
shoving lhe righl-handed allern, 3 er cenl shoving
no signihcanl asymmelry, and 16 er cenl shoving lhe
lefl-handed allern.
These frequencies of arm-lenglh asymmelry vere
almosl idenlical lo lhose recorded by Schullz (1937) in
analomy colleclions in lhe US, vhere lhe ercenlages
of inslances falling inlo each of lhe same calegories
vere 8O:4:16 in a ooled samle of 232 Americans of
Luroean anceslry. Schullz (1937) recorded dala on
long-bone lenglh asymmelries for lhe humerus, radius,
and for bolh combined, arlilioned by sex and also by
oulalion (his samle also included 233 Americans
of African origin, 122 Alaskan Lskimo-Inuil, 118 Norlh
American Indians, and smaller samles of Chinese and
of Aboriginal Auslralians). The overall incidence for lhe
vhole ooled samle is 79 er cenl longer righl arms,
3 er cenl equal lo measuremenl recision, and 18 er
cenl longer lefl arms. Ior all lhe oulalions for vhich
sex informalion vas labulaled, lhe females vere alvays
less likely lo have longer lefl arms and more likely lo
have longer righl arms (vhich is consislenl vilh sexual
dimorhism in lhe incidences of righl- and lefl-hand
reference: cf. Seddon & McManus 1991).
Moving back slighlly furlher in lime, ve can also
analyze skelelal samles from earlier oulalions of
modern humans (Hcmc scpicns scpicns). Thould &
Thould (1983) examined 416 adull skelelons from
Romano-rilish Ioundbury, and found lhal lhe arm
bones vere longer on lhe righl side in 21O individu-
als and on lhe lefl in 65 (lhe resl vere nol measurably
asymmelrical). A sludy of asymmelries in lhe radii in
a samle of 27 individuals from lhree Neolilhic farm-
ing siles in lhe Middle Llbe-Saale region, Germany,
found a righl-dominanl allern in 7O er cenl of
individuals, vilh 15 er cenl lefl-handed and 15 er
cenl 'ambidexlrous' (Reichel ci c|. 199O). Hovever,
lhe discriminanl funclion used lo redicl handed-
ness in lhis sludy is likely lo have somevhal inaled
lhe eslimaled frequencies of non-righl handedness.
In foraging eoles of early Holocene (Mesolilhic)
norlhern Luroe, mosl individuals sludied had longer
righl forelimbs, a allern seen slighlly more slrongly
in females (Conslandse-Weslermann & Nevell 1989).
24 adull males had longer righl arms (summed lenglhs
of lhe humerus and radius), and 9 had longer lefl arms.
Ior lhe females, lhe ralio vas 19 vilh longer righl
arms lo 5 vilh longer lefl arms.
Moving signihcanlly furlher back inlo evolulion-
ary lime, fossil hominin remains can also be analyzed.
There is only a very small number of individuals of
223
Humans, Tools and Handedness
exlincl secies vhose skelelons are reserved in suf-
hcienl comleleness lo enable leflrighl comarisons
of aired uer-limb elemenls. This limiled evidence
suggesls, hovever, lhal redominanl righl-handed-
ness exlends back in lime lo al leasl lhe early members
of our ovn genus Hcmc, around 1.6 million years ago
(mya). The skelelon of lhe Turkana boy from Nari-
okolome, WT-15OOO (early African Hcmc crccius, also
called Hcmc crgcsicr), has grealer develomenl of lhe
clavicular area of allachmenl of lhe righl delloid mus-
cle and grealer lenglh of lhe righl ulna, consislenl vilh
righl-handedness (Walker & Leakey 1993). Asymmelry
in lhe shafls of lhe humerus consislenl vilh righl-arm
dominance is also revalenl in Neanderlhal skelelons:
of six skelelons in vhich lhe relevanl measuremenls
could be laken bilalerally, all vere more robusl in
lhe righl arm (Trinkaus ci c|. 1994). The Neanderlhal
individual buried al Le Rgourdou, daled lo belveen
75 kya and 6O kya, also shovs several markers for
righl-handedness, such as a lhicker and more curved
righl clavicle, ulna, radius, and humerus (Vander-
meersch & Trinkaus 1995).
MatcrIa! cu!tura! markcrs nI handcdncss
The skelelal evidence is very sarse for Ileislocene
and earlier hominins. Technology rovides anolher,
more abundanl dala source. Archaeological evidence
from lools and olher arlefacls can also be used lo
infer lhe evolulion of human handedness, and is sum-
marized hrsl by melhod of eslimaling handedness
(sub|eclive hl in lhe hand, lool roduclion including
mullile ake analyses and knaing geslures, laler-
alized relouch, asymmelrical lool use and use-vear,
leelh marks, and arl including reresenlalions of lool
use, engravings, cave ainlings, and hand rinls), and
lhen vilhin each melhod, by lime eriod (from earliesl
lo mosl recenl evidence) and by malerial (slone, bone,
vood, anller, bronze, elc.).
Sub]ecIlte and earI assessnenIs
A number of early archaeologisls involved in excava-
lions have observed lhal cerlain lools hl beller in lhe
righl or lefl hand. Novadays archaeologisls refrain
from making such commenls because lhey are seen
as unscienlihc, bul in lhe lasl cenlury lhey vere ac-
celable. Ior examle, Gabriel de Morlillel (189O)
claimed lhal lhere vere more lefl-handers in rehis-
loric limes, based on Neolilhic double-edged scra-
ers from Irance and Svilzerland. Slrangely, he had
reviously (1883) argued for lhe oosile, hnding lhal
mosl 'hand-slones' of 'very early lribes' found in lhe
Somme gravels vere made for righl-hand use (ciled
in rinlon 1896). lher righl-hand suorling declara-
lions vere also made by lack ci c|. (1933), by Lvans
(1897) aboul handles and hafls for bronze sickles and
svords in Sviss lake dvellings and Lnglish barrovs
(see also Wilson 1891, 138), and by Sarasin ci c|. (ciled
in Senneman 1985 and Iosnansky 1959).
Such sub|eclive observalions vere based on an
inluilive suosilion of hov lo hold lhe lool, because
lhe gris, uroses, and manners of lool use vere nol
knovn. Nonelheless, il is inleresling lhal searale
researchers have made similar |udgemenls of male-
rial from vaslly dislanl siles bolh in lime and sace.
These early asserlions are lherefore vorlh including
in lhis reviev and may be vorlh revisiling in lhe fu-
lure, esecially nov lhal lhe melhods of griing and
using lools are becoming beller-knovn. Semenov's
(1964) volume is a good examle of lhe level of delail
lhal can be oblained in a sludy of use-vear in order
lo secify lhe recise kinds of hand conhguralion
lhal vere used lo gri lools during lheir use. Also,
recenl aers by Takeoka, Ihillison, and Iosnansky,
vhich vill be discussed belov, do lake gri osilion
inlo accounl.
Anolher early argumenl vas roosed by Darl
(1949), allhough his ideas are nov considered fanci-
ful. He hyolhesized lhal baboons from Slerkfonlein
had been hunled by lool-vielding hominins, as lhe
crania seemed lo shov signs of crushing from hand-
held bone veaons. These allerns, Darl suggesled,
indicaled redominanl righl-handedness because
a righl-hander holding a lool vill lend lo slrike lhe
fronl lefl (if from a face-on allack), or rear righl (if
from a sleallhy rear allack), arls of lhe viclim's skull.
Allhough Darl's hyolhesis vas develoed vilh lhe
scienlihc reasoning of his lime, our currenl knovledge
of lahonomy, as vell as conlinuing excavalions al
Slerkfonlein, has enabled crilicisms of Darl's ideas. Ior
examle, rain (1981, 2634, 1994) describes lhe lyes
of lahonomic rocesses involving carnivore gnav-
ing and roof collases lhal can roduce lhe damage
allerns observed by Darl.
TooI roducIlon
Tvo sludies have analyzed large assemblages of
akes lo hnd roorlions of righl and lefl akes,
lhey are lhose of radley & Samson (1986), and
Tolh (1985). In addilion, four sludies look al scal-
lers from single knaing evenls. Iischer (199O),
Hogberg (1999), Nevcomer & Sieveking (198O), and
Wenban-Smilh (1997) resenl dala from exerimenls
and archaeological siles shoving lhal lhe knaer's
handedness roduces a dislincl scaller allern on
lhe ground. The knaing geslure also imoses
conslrainls on accuracy and, lherefore, on lhe ho-
mogeneily of ake-surface allribules. Three sludies
224
Claplor 15
have used argumenls from lhe knaing geslure lo
indicale asymmelrical fealures on lools and akes
(Rugg & Mullane (2OO1), Takeoka (1991), and While
(1998)). Iinally, ve include an anecdolal reorl of a
lefl-handed knaer vho vas buried vilh his core
and hammerslone in hand.

TooI roducIlon nuIIlIe-fake anaIsls
An inuenlial sludy by Tolh (1985) roosed lhal
righl-handedness can be seen in lhe archaeological
record by reconslrucling lhe referenlial direclion of
core rolalion during inilial aking. The direclion of
rolalion of lhe core vas inferred from lhe resence of
corlex (lhe ouler surface of a inl nodule) on lhe righl
or lefl side of lhe dorsal surface of a ake. He sludied
akes from Koobi Iora (Kenya) from a number of siles
daled lo belveen 1.9 and 1.4 mya, redominanlly of lhe
ldovan induslry, bul included one Larly Acheulean
sile (daled lo belveen 1.4 mya and 7OO kya). Il is im-
orlanl lo nole lhal Tolh's melhod only alies lo a
secihc reduclion slralegy, namely lhe use of single-
lalform cores. This involves removing all lhe akes
from lhe same lalform, in sequence. n a round
cobble, lhis reduces lhe number of ossible aking
localions lo lvo: in fronl of lhe revious removal, or
behind il. Tolh's ovn relicalions of Karari scraers
roduced 56 er cenl righl-biased akes, caused by
rolaling lhe core clockvise in lhe lefl hand. He argues
lhal lhis decision is diclaled by 'lhe musculo-skelelal
slruclure of lhe lefl hand and arm, in vhich lhe su-
erior over of lhe suinalors and exors roduce
a referenlial rolalion in lhis direclion for a slronger
and more conlrolled lurning molion (. Love|oy ers.
comm.)' (Tolh 1985, 611). The hnding of 57 er cenl
righl-orienled akes al six Koobi Iora siles suggesled
lhal lhe Koobi Iora knaers, hominins from 1.6 mya,
vere al leasl as righl-handed as Tolh. He also sludied
Acheulean akes from Ambrona (Sain), daled lo
4OO3OO kya, and found 31 lefl-orienled for 48 righl-
orienled akes, a R:L ralio of 61/:39/.
In a sludy of Acheulean handaxes from Cadding-
lon, UK (daled lo 11513O kya), radley & Samson
(1986) relicaled biface and Levallois reduclion se-
quences by a righl-handed knaer. They classihed
lhe akes vilh resecl lo corlex relenlion as vell as
lhe resence and localion of relicl margins. A lenla-
live analylical melhod vas crealed vhich yielded a
handedness index of 62 er cenl R for lhe exerimenlal
colleclion, and 54 er cenl R for lhe archaeological
samle. The aulhors inlerrel lhese resulls as a veaker
bias lovards righl-handedness in Caddinglon com-
ared lo lhe exerimenlal knaing.
radley & Samson's classihcalion according lo
relicl margin localion means lhal reduclion sequences
are laken inlo accounl, a nolion vhich also aears in
Tolh's argumenl allhough il is oen lo crilicism. This
has lo do vilh exlending lhe Tolh melhod lo olher
archaeological colleclions. In facl, for mosl lyes of
knaing, lhe order of ake delachmenl is moslly
conlingenl on lhe shae of lhe core or inl nodule
(Iallerson & Sollberger 1986, Iobiner 1999). The
facl lhal lhe Karari cores vere aked from a single
lalform cerlainly alloved good serial aking. This
vas demonslraled by Ludvig & Harris (1994), vho
conhrmed lhal righl-handers rolaled lhe core clock-
vise and lefl-handers counlerclockvise vhen making
Karari scraers. Therefore ve musl be caulious vhen
alying Tolh's melhod lo induslries vhose reduclion
slralegies vere nol reslricled lo serial aking. Wilh
olher kinds of ake roduclion, lhe hgures seem lo
aroach 5O:5O as lhe samle sizes increase (Noble &
Davidson (1996), 17O, Iobiner 1999, Uomini 2OO1).
TooI roducIlon knalng scaIIers
Allhough lhere are very fev high-resolulion siles vilh
in siiu knaing scallers (e.g. Hogberg 1999), lhey are
valuable because lhey can reveal handedness. Il has
been shovn exerimenlally lhal a knaer silling
on a seal roduces a cenlral concenlralion of debris
vhich is skeved lo lhe side of lhe knaing hand.
Ior examle, Iischer (199O) describes a series of
con|oined arlefacls found in lace al lhe Trollesgave
sile, Denmark, near a large slone. The sile is daled lo
91OO and lhe arlefacls are referred lo lhe romme
lechnocomlex. Iischer, a righl-hander, exerimen-
lally relicaled lhe Trollesgave blades vhile silling on
a similar slone seal, in lhe evenl roducing a scaller
vhich vas mosl dense in fronl of his feel, fading oul
lo lhe sides, and righl-orienled. The archaeological
scaller vas also orienlaled lo lhe righl, localed simi-
larly in fronl of lhe slone. When silling direclly on
lhe ground, vilh one leg folded and one leg slraighl
oul, a clear lriangular scaller aears. Nevcomer &
Sieveking (198O) relicaled 16 Neolilhic axe roughouls
al lhe sile of Grime's Graves, UK. The lefl-hander,
Nevcomer, silling on lhe ground vilh his righl knee
benl and lefl leg oul, roduced a righl-skeved scal-
ler ending abrully vhere lhe legs vere. Il musl be
noled lhal lhere is anolher aclion vhich can roduce
a concenlraled scaller: lhe use of a iece of hide or
clolh for leg and crolch roleclion. The ieces vhich
fall on lhe malerial collecl inlo a dislincl hea vhen
lhey are dumed onlo lhe ground, for examle vhen
emlying lhe debris or vhen lhe erson slands u.
Il is imorlanl lo nole lhal lhe dumed hea looks
idenlical vhelher lhe knaing is done silling on lhe
ground, on a seal, or squalling (Nevcomer & Sievek-
ing 198O). Secihcally,
225
Humans, Tools and Handedness
lhe manner in vhich lhe roughoul vas held during
aking did nol seem lo have much effecl on lhe size
and shae of ake scallers. When lhe |sheeskinj
lhigh ad is used, il calches lhe akes as lhey are
slruck and lhe akes lhen lend lo dro in a circular
hea belov. Wilhoul lhe ad, vhich is difhcull lo
use vhen slanding or sealed on lhe oor, lhe akes
are eilher caughl in lhe hngers and lhen droed, or
alloved lo shool off freely, in eilher case lhe roughly
circular shae of lhe scaller is recognisable. (Nev-
comer & Sieveking 198O, 35O)
Similarly, Wenban-Smilh (1997), a righl-hander, sal
on lhe ground vilh lhe lefl leg folded and righl leg
oul (lhe inverse of Nevcomer) and roduced a lefl-
skeved scaller bounded by lhe legs.
TooI roducIlon knalng gesIure
Wilh resecl lo knaing geslures, Takeoka (1991)
dehnes lvo kinds of movemenl vhich affecl lhe o-
silion of lhe ake blank (or core), and lhus lhe angle
al vhich il receives lhe hammerslone blovs. ne is
vrisl abduclion1adduclion, lhe olher is forearm ro-
nalion1suinalion. When knaing, lhe axis of vrisl
movemenl (if lhe alm is laced al on a lable, lhis
vould be a side-lo-side molion of lhe hand) affecls
lhe direclion of fraclure force roagalion vilhin
lhe core, lhis is lhe effecl lhal lhe cone of ercussion
melhod exloils, allhough lhey argue for an enlirely
hammerslone-based cause (Rugg & Mullane 2OO1).
Iorearm rolalion affecls lhe vorking angle (angle
belveen lhe lalform and hammerslone lra|eclory),
a more ronaled vrisl resulls in an obluse angle (be-
cause lhe lalform is lilled lovards lhe body) vhile a
more suinaled vrisl resulls in an acule angle (lal-
form lilled avay from lhe body). A lhird faclor, vrisl
exion1exlension, affecls lhe horizonlal osilion of lhe
slriking lalform, bringing il closer lo lhe knaer's
eyes (Takeoka 1991, 5O35).
Rugg & Mullane hyolhesize lhal:
lhe angle al vhich lhe cone of ercussion occurs
relalive lo lhe slriking lalform is usually around
9O degrees, bul can vary ... ecause lhe human arm
has ivol oinls al lhe shoulder, elbov and vrisl, il
is lausible lhal some blovs vould lead lo cones of
ercussion lhal vere angled lo lhe righl or lefl relalive
lo lhe slriking lalform. (Rugg & Mullane 2OO1, 252)
ecause lhe Herlzian cone indicales direclionalily, ils
skev should reecl lhe exacl lra|eclory of lhe hammer-
slone. Rugg & Mullane exerimenlally validaled lheir
recognilion crileria, vilh four lefl-handed knaers
and four righl-handers: in a blind lesl lhey vere able
lo assign 75 er cenl of lhose akes lhal had a clear
cone of ercussion lo lhe correcl handedness.
The facl lhal righl-handers roduced righl-
skeved cones and lefl-handers roduced lefl-skeved
ones indicales lhal lhe lendency lo skev lhe blov
comes from eilher slighl, uninlended suinalion of
vrisl or uninlended exion al lhe elbov of lhe kna-
ing arm. The basic knaing geslure, as described
above, consisls of arlially ronaling lhe vrisl and
simullaneously adducling lhe forearm, so any devia-
lion lo orienl lhe blov lovards one's body is caused by
exlra suinalion and1or exion. These biomechanical
suggeslions deend on lhe bimanual conhguralion
used in knaing, vhich is discussed belov.
Ior simlifying uroses, ve vill say lhal kna-
ing can be done vilh hve general hand osilions,
or conhguralions. The hrsl four involve holding lhe
core againsl one leg and are groued as lvo differenl
lechniques: Ilake Suorl and Iree Iall. We suggesl
lhese names lo reecl lhe immediale inlenlion for
lhe resulling akes. Nevcomer & Sieveking (198O)
refer lo 'Iree Iall' vs 'Deliberale lacing in a hea',
lo dislinguish vays of lrealing blades as lhey come
off lhe core. Generally, Ilake Suorl is used vhen
lhe ake ilself is lhe inlended roducl, meaning lhe
knaer vanls lo revenl il from falling lo lhe ground
vhere il mighl break, lhe core is ressed againsl lhe
lhigh so lhal lhe resulling ake vill lie sandviched
belveen lhe core and leg. In Ilake Suorl, lhe core is
held eilher againsl lhe ouler surface of lhe isilaleral
(same side as core hand) lhigh, or lhe inner surface of
lhe conlralaleral lhigh. Conversely, Iree Iall lends lo
be used vhen lhe akes are vasle roducls, in lhis
case, lhe core is ressed againsl lhe leg so lhal lhe
ake comes off lhe 'free' side of lhe core and falls lo
lhe ground. In Iree Iall, lhe core is held eilher againsl
lhe inner side of lhe isilaleral lhigh (aking belveen
lhe legs), or lhe ouler side of lhe conlralaleral lhigh
(aking on lhe oulside of lhe body, vhere lhe kna-
ing arm has lols of sace lo move).
Hovever, lhese suggeslions are by no means slricl
rules, as lhe shae of lhe ake is diclaled by lhe vay lhe
hammer's energy is lransferred. Secihcally, in Ilake
Suorl, lhe hammer arm's lra|eclory is sloed by lhe
leg, causing lhe energy lo ov inlo lhe leg, lhis lends
lo roduce curved lhinning akes. In Iree Iall, lhe
hammer arm can follov lhrough ils lra|eclory, resulling
in lhe energy going lhrough lhe core, roducing al
lhinning akes (. radley ers. comm.).
An imorlanl faclor affecling knaing conhgu-
ralion is lhe lechnique, dehned by lhe lye of hammer
(I. Slernke ers. comm.). A sofl hammer (i.e. anller)
requires much grealer velocily, meaning one lends
lo kna vilh Iree Iall so as nol lo smash one's leg.
Hard hammers (i.e. mosl slones) can be vielded vilh
less seed, so il is ossible lo use Ilake Suorl. Also,
vhen lhinning a handaxe, lhe core is normally held
in lhe hand or vilh Ilake Suorl. This gives more
226
Claplor 15
conlrol of core salial conhguralion, and revenls end
shock (an unrediclable accidenl causing lhe handaxe
lo sna in lvo).
Addilionally, one can enlirely suorl lhe core in
one hand (Nevcomer & Sieveking call lhis Ireehand:
198O, 349). Ior large or heavy ob|ecls lhe forearm can
be suorled in lurn by lhe isilaleral lhigh (lhis
conhguralion is observed in knaers from Iaua
Nev Guinea: see Sloul 2OO2). Ior small ob|ecls or
vhen hner conlrol is needed, lhe core is held in lhe
unsuorled hand, such as in lhe lhinning slage of
handaxe roduclion.
In addilion lo lhe hve menlioned above, olher
conhguralions do exisl, such as holding lhe core be-
lveen lhe lvo legs (as in indirecl ercussion for blade
roduclion), or holding lhe core againsl an ob|ecl
(anvil, lree slum, ground, elc.). Il is also imorlanl
lo nole lhal some melhods (core-reduclion slralegies)
make use of more lhan one knaing conhguralion.
Ior examle, making blades by direcl ercussion hrsl
requires making a cresl, vhich 'is done on lhe oulside
of lhe lhigh' (Nevcomer & Sieveking 198O, 35O), and
second, lalform rearalion folloved by blade re-
movals, bolh 'done belveen lhe legs' (Nevcomer &
Sieveking 198O). In his blade exerimenls, Nevcomer
roduced lvo dislincl scallers searaled by his leg,
conlaining lvo differenl lyes of debilage, reecling
lhe use of lhese lvo conhguralions.
The manner of holding lhe core can also inleracl
vilh lhe reduclion slralegy, indicaling handedness.
While (1998) idenlihed four ossible bimanual con-
hguralions for manufacluring lvisled ovales. These
bifaces exisl in rilish siles daled lo from lale IS-11
lo early IS-1O in signihcanl roorlions (2O46 er
cenl) al siles like ovman's Lodge, Wansunl Iil, and
Svanscombe (all lhree in Kenl), Llveden and Ioxhall
Road (Suffolk), Allinglon Hill (Cambridgeshire), and
Hilchin Lake eds (Herlfordshire), and in Irance dal-
ed from IS-12111 lo ossibly IS-8. Tvisled ovales
are made vilh a arlicular melhod, usually al lhe
hnishing slage: hrsl, one quarler of lhe edge is aked
unifacially. Then lhe handaxe is inverled lhrough lhe
long axis and one quarler of lhe oosile face is aked.
These lvo sels of unifacial removals, on oosing
faces, are nov |oined al one li of lhe handaxe. Nexl,
lhe iece is rolaled (clockvise or counlerclockvise)
18O degrees and one more quarler aked unifacially.
Iinally, lhe iece is inverled lhrough lhe long axis
again and lhe oosile quarler is aked, bringing lhe
lasl lvo sels of removals lo |oin al lhe olher end of lhe
handaxe. The resull is a handaxe vilh an edge aller-
naling four limes belveen lhe lvo faces. This makes
lhe rohle look 'lvisled' in lhe same vay, no maller
hov you hold il.
Ior all four edges lhal are knaed unifacially,
il is lhe handaxe vhich is rolaled so lhal lhe ham-
mer hand alvays knas in lhe same 'aclive zone' of
lhe core hand (While 1998, 99). The inlerrelalion of
handedness comes from lhe facl lhal nearly all lvisled
ovales have a Z-shaed rohle ralher lhan an S shae.
This means lhal lhere are lvo ossibililies for lhe
aclive zone: eilher lhe area near lhe vrisl for a righl-
hander, or lhe area near lhe hngers for a lefl-hander.
(A righl-hander using lhe hngers area, as vell as a
lefl-hander using lhe vrisl area, vould roduce an S
lvisl.) The use of lhe hngers area can only be |uslihed
if lhe rehisloric knaers vere moslly lefl-handed,
and so lhis ossibilily can be excluded, leaving only
lhe righl-handed olion as an exlanalion of lhe
Z-shaed rohles.
Iinally, a remarkable burial vas found al Hazel-
lon Norlh, Colsvolds, UK, daled lo 55OO (Saville
2OO3). This lomb conlained a male, 3O45 years old,
vilh a inl core benealh his righl elbov, and a ham-
merslone al lhe lace of his lefl hand. The core and
hammer vere mosl likely laced inlo lhe burial afler
his dealh, lhey vere eilher laced failhfully, meaning
he vas a lefl-handed knaer, or he vas righl-handed
and lhey vere laced incorreclly, meaning his buriers
disregarded lhe hand he used vhen knaing.
LaIeraIl:ed reIouch
Iive aulhors (Cornford 1986, Ihillison 2OOO, Semenov
1964, lankholm 199O, rinlon 1896) describe evidence
of handedness from asymmelrically-relouched lools.
This asymmelry can be due lo laleralized use, making
il necessary lo relouch lhe more vorn side of lhe lool,
or simly from conslrainls in knaing vhen holding
lhe iece. Cornford (1986) describes akes resulling
from a ccup !c ircncnci. The sile of La Colle de Sl
relade, Irance has a long slraligrahy sanning lhe
lasl lvo inlerglacials (from 24O kya lo 122 kya). The
lools vere resharened vilh a lranchel blov lo freshen
one edge of one face of lhe li. These sharening akes
vere removed by righl-handers in roorlions ranging
from 77 er cenl lo 91 er cenl, from oldesl lo young-
esl layers of lhe sile. Iurlher evidence comes from
lhe use of bone relouchers (Semenov 1964, 163). The
arlefacls come from Middle Ialaeolilhic (Kiik-Koba
and Teshik-Tash) lo Uer Ialaeolilhic (Koslenki 1)
siles in Russia, daled lo 3734 kya. Semenov indicales
denls on lhe convex side of bone relouchers vhich
mel al an angle of 7585 degrees lo lhe long axis, sug-
gesling lhey vere used by righl-handers. lankholm
(199O) also found laleralized relouch on microlilhic
armalures of lhe Maglemosian induslry (9.58 kyr
) in soulhern Scandinavia. These lanceolales and
lriangles vere moslly relouched on lhe lefl side, in
227
Humans, Tools and Handedness
roorlions from 5O er cenl lo 1OO er cenl, and lhis
vas nol due lo any funclional conslrainls. Ihillison
(2OOO and ers. comm.) noles a sile conlaining cherl
scraers al Aksum, Llhioia, daled lo lhe hflh lo sixlh
cenluries , vhich are asymmelrical in shae. They
consislenlly have one surred corner, usually on lhe
lefl, and vhich Ihillison allribules lo habilual use
in lhe righl hand. An early sludy of Norlh American
Indian lools by rinlon (1896) measured lhree asym-
melrical crileria: offsel oinl asymmelry, side of laleral
relouch, and blade 'lvisling'. He found aboul lvo-
lhirds more righl-handed fealures lhan lefl-handed
ones in undaled blades of cherl and |aser (hio),
cherl, quarlz, and |aser (Wisconsin), and argillile,
|aser, quarlz, and black cherl (Nev }ersey).
Use and use-uear
Tvo aulhors examined lhe lraces lefl on slone lools
by use, secihcally scraers and akes (Irame 1986,
Semenov 1964). Three aulhors include secihc con-
slrainls aboul gris in lheir analysis (Takeoka 1991,
Ihillison 1997, Iosnansky 1959). Tvo aulhors and
colleagues sludied use-vear from a rolaling molion
(Keeley 1977, Cahen ci c|. 1979, Cahen & Keeley 198O,
Senneman 1987). Roosevell (1974) describes asym-
melrical vear on vooden soons. Gerharz & Sen-
neman (1985), along vilh Wilson (1885, 1886, 1891),
reorl evidence of use-vear and use allerns from
bronze sickles. Three aulhors oinl lo lhe necessily of
hiring lefl-handed miners in Roman limes.
The La Colle arlefacls vere examined for mi-
croscoic use-vear lraces lo delermine handedness
by anolher lechnique. Irame (1986, in lhe Cornford
volume) insecled lhe slrialion orienlalion and bands
of olish from vorking vood, hide, or olher malerials
on long sharening akes. f 18 righl-asymmelrical
sharening akes, 4 had oblique righlvard marks, 1
lefl, and lhe remainder eilher erendicular, arallel,
or mullidireclional. f 4 lefl-asymmelrical akes, 2
had lraces of moving leflvard, 1 erendicular, and 1
mullidireclional. Irame roosed lhal lhese marks, in
relalion lo lhe vorking edge indicaling lhe direclion
of lool use, shoved lhey vere referenlially used by
righl-handers.
Semenov (1964, 87f.), in his volume on use-vear,
described lhe mechanism for asymmelrical scraer
vear. The scraers vere used on hide, vilhoul han-
dles, simly held in lhe hand. ecause lhe lool is
held 'vilh ils axis al an angle of 758O degrees lo lhe
skin surface', by imlicalion, lhere is a conslrainl on
simullaneous abduclion of lhe uer arm and rona-
lion of lhe vrisl1forearm (in orienling lhe lool-using
hand erendicular lo lhe surface being vorked). This
imlies lhal force is more efhcienlly exerled vhen lhe
arm is less abducled and lhe forearm less ronaled. Se-
menov counled lhal aboul 8O er cenl of end-scraers
are vorn on lhe righl side. His dala include Russian
(Koslenki 1, Timonovka, Mezin, Suonevo, Saka|ia) as
vell as olher Uer Ialaeolilhic siles. Takeoka (1991)
furlher argued from scraer usage, vilh lhe assum-
lion lhal lhe scraer vas ulled lovards one's body,
lhe venlral surface al lhe fronl. In lhis molion, lhe
lhumb is ressed againsl lhe venlral surface, hngers
suorling lhe dorsal surface. Takeoka argued lhal
lhe vorking edges of lhe scraers are moslly localed
on lhe side of lhe ake lhal vill ul lhe roximal
(lhickesl) end of lhe ake inside lhe cued alm,
ralher lhan lhe hngers, and lherefore vere made for
righl-handers. Ihillison (1997) conhrmed lhis effecl.
She scrulinized 54 handaxes and cleavers recovered
by an LS Leakey excavalion in 1931 in Kenya. Their
slraligrahy is daled lo aboul 1 mya. Slarling from
lhe remise lhal lhe lrailing face, nol lhe leading
face, of a used edge, vould shov grealer signs of use,
Ihillison reconslrucled ossible gri lyes for each
iece. f 54 lools, 6 (11 er cenl) could be assigned
lo robable lefl-hand use, 45 lo lhe righl hand, and
3 vere indelerminale. Conslrainls of use involve lhe
efhcienl exerlion of force and resislance of hnger and
hand muscles:
rolalion of lhe vrisl vilhoul shifling hngers ermils
lhe concave edge lo be used as a ull scraer. (Ihil-
lison 1997, 18O)
Some imlemenls had more lhan one vorking edge, and
so by imlicalion vere held in several differenl vays:
These osilions vould have alloved for a number of
lyes of force lo be exerled in several direclions vilh
lhe lool, deending uon exaclly hov lhe hand vas
laced. A line of force from lhe vorking edge lhrough
lhe cenlral mass of lhe lool lo lhe base of lhe alm of
lhe hand, for examle, ermils sleady ressure lo be
alied vhile lhe hngers are arlially freed lo rolale
lhe lool in sublle scooing, lvisling or scraing mo-
lions. A gras in vhich lhe handaxe is comressed
belveen lhe lis of lhe hngers and lhe alm of lhe
hand is needed lo revenl loss of conlrol vhen il is
used for heavy culling or saving in a direclion ar-
allel lo lhe ulilized edge. The shock of choing or
digging molions is besl absorbed by lhe fronl of lhe
alm of lhe hand or lhe base of lhe hngers, allhough
a oslure vilh lhe hngers vell sread and lhe force
falling somevhal furlher forvard is also effeclive.
(Ihillison 1997, 174)
Iurlhermore, an asymmelrical veighl dislribulion on
lhe lool can facililale use:
a hand-hold vas rovided by a relained area of
lhe original corlex or a ake slriking lalform on
an olhervise bifacially vorked secimen. In mosl
inslances lhis more rounded area vas associaled
228
Claplor 15
vilh an asymmelric bulge on one or bolh faces of lhe
handaxe vhich hl comforlably inlo lhe concavily of
lhe user's gras and greally facililaled lhe conlrolled
maniulalion of lhe lool. (Ihillison 1997, 174)
This laller slalemenl, like lhe nexl one belov, is an
examle of hov lo reconcile lhe nineleenlh-cenlury
sub|eclive observalions menlioned above vilh lhe
rigorous scienlihc aroach referred loday. A similar
observalion on lhe use-conslraining effecls of asym-
melrical veighl dislribulion in lhe arlefacl vas made
by Iosnansky (1959), in sludying a colleclion of Larly
lo Middle Acheulean handaxes from lhe Trenl Valley
(UK) and 118 handaxes from lhe Iurze Ilall sile (UK).
He slales:
il is found lhal lhe dislacemenl of lhe veighl avay
from lhe culling edge, vhich a non-cenlral median
ridge imlies, increases lhe efhciency for culling.
(Iosnansky 1959, 42)
Like Ihillison, Iosnansky lesled lhe handaxes for
ease of use in eilher hand, assuming a culling funclion.
Secihcally, 'lhe mosl efhcienl melhod of culling is
one in vhich lhe bull of lhe lool is held in lhe alm of
lhe hand vilh lhe hngers slayed around lhe blunler
of lhe lvo edges and lhe al face of lhe lool faces lhe
inner cul face' (Iosnansky 1959, 43). f 4O comlele
lools in lhe Turlon colleclion, 35 er cenl vere found
lo beller accommodale lhe righl hand, 12.5 er cenl
lhe lefl hand, and 52.5 er cenl eilher hand. Tvo in-
deendenl observers found similar roorlions: lhey
assigned lhe folloving reseclive roorlions lo lhe
same handaxes 37.5 er cenl-15 er cenl-47.5 er cenl
and 22.5 er cenl-1O er cenl-67.5 er cenl.
Keeley (1977) describes a biface from Claclon
(2OO kya) vilh microscoic use-vear shoving il vas
used vilh a verlical rolaling molion, such as boring
holes, in a clockvise direclion. Keeley's argumenl
imlies lhal grealer lorque forces are exerled during
vrisl suinalion (clockvise for a righl-hander) lhan
ronalion. Indeed, suinalion roduces more lorque
lhan ronalion (Sellers 2OO4), and lhis is lhe reasoning
behind lhe design of screvs: lhey musl be screved
in clockvise, vhich exloils lhe slronger suinaling
lorque of lhe righl hand. The mode of rehension is
nol secihed, bul a lool being verlically rolaled can
be held eilher vilh lhe elbov u and alm facing
oulvard (screvdriver gri), or vilh lhe elbov dovn
and alm invard (slabbing gri). This resuoses
lhal vhalever lhe gri on lhe lool, eole grind in a
direclion oulvard from lhe cenlre. In a screvdriver
gri, lhe vrisl musl roduce mainly suinaling
forces, vhile grinding vilh a slabbing gri, lhe vrisl
roduces mainly exlensor forces. olh of lhese could
reecl a reference lo suinaling1exlension ralher
lhan ronalion1exion (vhich vould be lhe forces
required if lhe grinding molion venl invard). Cahen
ci c|. (1979) conhrm lhis conslrainl:
Allhough a back-and-forlh lurning of lhe borer is
efhcienl vhen lhe borer is hand-held, lhe oulvard
lurn of lhe vrisl is more overful. Lxerimenlal
observalions have shovn lhal lhe relurn slroke in
lhe veaker, invard direclion is usually accomanied
by a slackening of lhe verlical ressure. (Cahen ci c|.
1979, 668).
In olher vords, boring is usually done vilh a back-
and-forlh molion, bul lhe oulvard slroke roduces lhe
bulk of lhe slrialions. In addilion, microvear olish
and edge damage indicale lhe rincial direclion of
lurning:
Generally seaking, microvear olish forms on lhe
asecl of edge ridges and ro|eclions facing lovard
lhe rincial direclion of lurning, vhile ulilizalion
damage is crealed mosl heavily (somelimes only)
on lhe asecl facing avay. (Cahen ci c|. 1979, 681, in
rely lo ob|eclions from Nevcomer and dell).
Using lhe same analysis melhod, Cahen and col-
leagues (1979, Cahen & Keeley 198O) sludied inl
lools from Meer. This elgian sile vas excavaled
in lhe 196Os and 7Os, and is daled lo 9 kyr . The
lilhic assemblage is characlerized by T|onger oinls,
vhich are backed blades used in ro|ecliles. They
examined lhe use-vear on 31 lools vhich had been
used for boring holes in, and engraving, bone and
anller. These lhick-billed borers are called becs. The
becs vere groued according lo vhich inl block
lhey could be rehlled lo, lhere vere 6 rehl grous in
lhis samle. 21 becs had been used clockvise and 3
counlerclockvise. These lhree becs vere all knaed
from lhe same block, suggesling lhey vere made and
used by a single erson. The aulhors conclude lhal lhe
main knaing scaller, called Concenlralion IV, vas
roduced by al leasl lvo eole, one of vhom vas
lefl-handed (Cahen ci c|. 1979, 671).
Anolher sludy using lhe marks from rolaling mo-
lions vas made by Senneman (1984a), vho examined
Sviss and German Neolilhic bone, anller, and slone
grinding lools. These lools dislay slrialions running
from lhe lo lefl lo bollom righl (for a righl-hander)
and lo righl lo bollom lefl for a lefl-hander. The siles
are all daled lo belveen 4O5O and 29OO , conlain 19.4
er cenl (of 31) lefl-handed lools al urgerrolh, Ger-
many, 19.6 er cenl (of 51) al odman, Germany and
6.3 er cenl (of 597) al Tvann, Svilzerland.
Roosevell (1974) examined a series of vooden
and bone soons and salulas from norlhern Chile.
They come from lhe Chiu Chiu sile and are daled lo
1OOO15OO. Roosevell also crealed an exerimenlal
sel of vooden soons used, righl-handedly, lo slir and
scoo food in a coarse ceramic bovl, and lhese shoved
229
Humans, Tools and Handedness
laleral vear on lhe dislal end. The archaeological
soons consisled of 76 delerminable lools, of vhich
48 vere righl vorn, 4 lefl vorn, and 26 bilalerally
vorn. ecause lhe archaeological and exerimenlal
soons shov idenlical vear allerns, lheir usage for
mixing food as vell as igmenl and snuff vas con-
hrmed. one and vooden salulas vere also robably
used vilh lhe concave arl facing lovard lhe user and
lhe handle slighlly angled (nol verlical), as lhis is lhe
mosl effeclive vay lo hold lhem (Roosevell 1974, 1O2).
This imlies arlicular conslrainls on combinalions
of molor acls vhen erforming a slirring or ladling
molion.
Senneman (1987, 22) offers some suggeslions
for idenlifying use-vear in adzes:
In general, lhe movemenl of a hafled adze blade is
nol verlical bul oblique in relalion lo lhe oeralor's
body. This is due lo lhe nalure of lhe ball |oinl be-
lveen lhe shoulder blade and lhe humerus. This
angle is more oblique if lhe adze is held in one hand,
lhan vhen held in bolh. In case of a righl-handed
individual, lhe movemenl runs from lo lefl lo bol-
lom righl (as vieved by lhe vorker). . Due lo lhis
slighlly oblique movemenl lhe edge of lhe slone
adze does nol hil lhe vorked malerial in an olimal
manner, one end of lhe edge making conlacl earlier
lhan lhe olher.
In alicalion of lhese conslrainls, Gerharz & Sen-
neman (1985) describe lvo bronze farming lools from
norlhern Llhioia shoving use by lefl-handers. The
lools can be relaled lyologically lo induslries around
1OOO , allhough direcl daling of lhe sile disagrees.
Several casl adzes from Yeha, a sile conlemorary vilh
Haoulli, have asymmelrical use-vear. The uer face
is more vorn on lhe righl corner, indicaling il vas
used by a lefl-hander. Anolher adze from lhe same
sile shovs righl-handed use, as do lvo olher adzes
from olher norlhern Llhioian siles. Iurlhermore, one
casl sickle from Haoulli, daled lo belveen 3OO and
1OO, is vorn from inlensive use. Il vas made for a
lefl-hander because il has a slrenglhening rib running
along lhe uer face, and lhe uer face vas hafled
on lhe lefl, meaning il vas held in lhe lefl hand. The
aulhors reorl hnding, among 8OOO bronze sickles,
only four olher lefl-handed sickles (lvo German ones,
one Hungarian and one Romanian).
The use of lefl-handed vorkers has been docu-
menled in Roman limes, in lhe conlexl of mining. Roder
(1957) describes lhe Roman mines for luff (consolidaled
ash) in lhe Iellenz, rohl Valley, Rheinland (Germany),
in vhich lhere vere usually 3 lefl-handers for 2 righl-
handers (3:2), or even 2:1.The vorkers used rods lo dig
oul lhe valls of lhe mines, making verlical valls. The
lefl valls vere slraighlened by righl-handers, and lhe
righl valls had lo be slraighlened by lefl-handers (e-
don 1984, 158). In lhe Gallo-Roman mine al Sainl-oil
(Irance, 1sl cenlury ), lhe reclangular blocks had lo
be carved oul by lvo miners vorking logelher, one of
each handedness (Monlhel 2OO2, 96). The biomechani-
cal conslrainls of vorking close lo a vall vilh a mining
rod meanl lhal lhe roorlions of righl and lefl handed
miners had lo be carefully selecled, lhese should be
visible in lhe vrillen records lhal some miners kel of
lheir vorkers (G. Monlhel ers. comm.).

CuI narks on IeeIh
Three sludies have sludied lhe marks lefl on Nean-
derlhals' anlerior denlilion by using slone lools lo cul
meal held belveen lhe leelh. This is elhnograhically
common (Semenov 1964, 1O4):
Generally asloral or hunling eole (like lhe
nomads of Mongolia, Tibel, Abysinnia and olher
counlries) eal such meal vilh a knife in one hand.
Meal is normally cul inlo slris, and baked or cured
in lhis form. Then each erson lakes a iece and,
holding one end in his leelh, culs il free vilh a quick
movemenl of lhe knife al his moulh, reealing lhe
oeralion unlil lhe vhole slri has been consumed.
The culling is done uvards from belov. We have
seen lhis done among Nenelz reindeer herdsmen in
lhe Kanin eninsula in 1928.
The slrialions on Neanderlhal and re-Neanderlhal
hominins' leelh vere examined by ermdez de Cas-
lro ci c|. (1988) and by Iox & Irayer (1997). In addilion,
lhe oxgrove hominid has similar marks on ils lvo
leelh. If one dislikes lhe idea of using lhe leelh as lools
(e.g. ax & Ungar 1999, vho exlicilly re|ecl a connec-
lion belveen handedness and slrialion orienlalion),
lhere is also lhe inleresling ossibilily lhal lhese marks
vere made by chiing inl vilh lhe leelh, an aclion
vhich has been observed in Ilains Indians (USA) and
Auslralian Aborigines (Hesler 1973).
ermdez de Caslro ci c|. (1988) reorled on
slrialions found on lhe fronl leelh of Hcmc nci!c||cr-
gcnsis individuals from lhe Sima de los Huesos sile
(Alauerca, Sain) (19 leelh, comrising 4 individuals
and 1O unassigned leelh), lhe La Quina 5 Neanderlhal
(2 leelh), one isolaled loolh from Cova Negra, Horlus
(several anlerior leelh from 5 individuals), and include
ublished dala on Sainl rais (1 isolaled loolh), An-
gles-sur-l'Anglin (1 isolaled loolh), and lhe Shanidar 2
Neanderlhal (2 leelh). Rough dales for lhese fossils are
as follovs: Alauerca aboul 3OO kya, La Quina 353O
kya (a Irench cave vilh arlefacls from a Mouslerian
area and an Aurignacian-Chlelerronian area), Cova
Negra 12O35 kya (a Sanish cave vilh many slone
lools, faunal, and Neanderlhal remains), Horlus 6O55
kya (a Irench cave vilh many Neanderlhal remains
23O
Claplor 15
and slone lools), Sl rais 5O4O kya (a Sviss cave),
Angles-sur-l'Anglin 1413 kya (a middle Magdale-
nian rock sheller vilh ainlings), Shanidar 6O kya (a
Neanderlhal cave burial in Iraq).
Ior comarison, a rognalhic moulh-guard
vilh fake enamel Neanderlhal leelh vas vorn by a
righl-hander. The exerimenlal rocedure involved
holding a iece of meal belveen lhe fronl leelh and
culling off bile-sized ieces vilh inl akes. The
exerimenler made slrialion allerns consislenl vilh
a righl-handed dovnvard molion from lefl lo righl
(vhen vieved from lhe fronl), malching lhose on lhe
fossil leelh. All of lhe fossil samles (excel Angles,
vhich has horizonlal marks, and Horlus VIII) shov
slrialions oinling dovnvard lo lhe righl. The leelh
from Horlus VIII has inversely orienled slrialions,
suggesling lhis individual vas a lefl-hander.
The aulhors slale lhal 'in lhe exerimenlal sludy,
lhe aclion lhal vould roduce slrialions as in scheme
C |consislenl vilh a righl-handed oeralor culling
leflvardsj vas uncomforlable and fell less efhcienl' (.
41O). This imlies lhal il vould be equally uncomforl-
able and inefhcienl for a lefl-hander lo cul righlvards,
and lherefore lhe observed righlvard fossil slrialions
reclude a lefl-handed roducl. The inefhciency of lhe
righl-handed leflvard molion imlies a conslrainl on
simullaneous ronalion al lhe vrisl and exlension of
lhe forearm. A righl-hander culling from belov (as
described by Semenov) vould roduce allern D on
his1her leelh, consislenl vilh a lefl-hander culling
leflvard from above. ul ermdez de Caslro ci c|.
(1988) did nol observe allern D in lhe fossils, sug-
gesling lhe culling vas done dovnvards by righl-
handers. Iurlhermore, lhe aulhors idenlihed arlial
Herlzian cones in lhe slrialions vhich indicale a
dovnvard culling direclion.
Iox & Irayer (1997) sludied lhe leelh of Kraina
Neanderlhals, vhich also shov slrialions consislenl
vilh culling meal held belveen lhe leelh. Six of lhe
lhirleen individuals above age lhirleen vere found
lo dislay righlvards scralches, vilh one shoving
leflvards scralches. The remaining six individuals
shoved no redominanl allern (|udged by lhe 5O
er cenl mark). Iox & Irayer also include ublished
dala from Kabve and Tabun individuals, vho also
have righl-handed slrialions. Iurlher evidence comes
from lhe lvo oxgrove (4OO kya) hominin leelh, vhich
bolh came from lhe same moulh and vere ad|acenl
bollom fronl leelh, and shov similar slrialions. They
also indicale righl-handed culling vilh inl (Iills &
Roberls 1997, 265).
In lolal, lhese lhree sludies reveal only 2 lefl-
handed hominins for 19 righl-handers. The number of
individuals of unknovn or indelerminale handedness
is 7 in lhese sludies, bul ve mighl assume lhal lhe
roorlion of righl lo lefl (1O.5 er cenl) is roughly
similar in lhe indelerminale samles.
ArI
Uer Ialaeolilhic and laler arl is a furlher source
of evidence for handedness. ne suggeslion is lhal
lhe ainlings and engravings are easier lo read, and
lherefore vere made, vilh a lighl source coming
from above on lhe lefl side (Delluc & Delluc 1993, 44),
meaning lhe arlisls held lheir lorch in lhe lefl hand
because lhey needed lo use lheir righl hand lo drav
and engrave. lher forms of evidence are images of
eole using lheir hands, engraved ebbles vhose
characlerislic lraces shov lhe direclion of engraving,
lhe draving of animal silhouelles, and lhe roorlions
of handrinls and hand slencils made vilh lhe righl
and lefl hands.
Rcprcscniciicns cj |cicrc|izc! icc| usc
Iive sludies have counled lhe number of deiclions
of righl- and lefl-handed lool use in vorks of arl.
Uhrbrock (1973) made an exlensive reviev of laler-
alily deicled in ainlings, scullures, medallions,
coins, and slams. He reorls higher roorlions of
lefl-facing rohles on US coins and medallions, bul
more righl-facing rohles on Luroean coins and
medallions daling from 6OO lo 1964. Iainled
orlrails are also slighlly more likely lo deicl righl-
facing eole. Deiclions of lool use in scullures,
columns, and dravings from lhe Renaissance lo
modern limes shov righl- and lefl-handedness, and
reresenlalions of lhe Madonna holding her Child
frequenlly shov her holding him on her lefl side
(Uhrbrock 1973). This mighl reecl eilher a conscious
choice of lhe ainler1scullor, or lhe need for righl-
handed molhers lo kee lheir dominanl hand free.
Anolher reviev vas made by Coren & Iorac (1977),
vho labulaled 118O inslances of unimanual lool and
veaon use deicled in dravings, ainlings, and
scullures from Luroe, Asia, Africa, and America,
sanning lhe lime 15,OOO lo 195O. 92.6 er cenl
of lhese images orlrayed righl-hand use, remaining
signihcanlly consislenl across geograhical areas
and lime eriods. lher such sludies are described
by Senneman (1984a, 613), Dennis (1958) sludied
Lgylian ainlings in lombs and found 7.5 er cenl
(n 12O) lefl-handed aclions in lhe 25OO -daled sam-
le, and 4.76 er cenl (n 191) lefl-handed deiclions
in lhe more recenl samle. An assessmenl of hand
use deiclions vas made by Senneman (1984b), for
a decoraled uddhisl yramid in Cenlral }ava vhich
vas conslrucled belveen lhe eighl and lenlh cenluries
. Senneman sludied 15O4 scenes (none deicling
231
Humans, Tools and Handedness
lhe uddha). These reliefs deicl 14,892 eole, 1O85
of vhich are using lheir hands unimanually. Mosl (926
83.5 er cenl) vere simly holding an ob|ecl, and
only 153 ( 16.52 er cenl) vere erforming a skilled
aclion. The unskilled aclions include leaning on a slick,
holding a horse, a svord, a over, axe, fan, umbrella,
or reaching for somelhing. The skilled aclions include
laying an inslrumenl, maniulaling food, riding an
elehanl, and using a veaon such as a svord, knife,
bov and arrov, or sear. The righl lo lefl ralio for
skilled aclions vas found lo be 137:16 (89.5 er cenl
righl-handed), vhile lhe ralio for unskilled aclions
vas 578:348 (62.4 er cenl righl). Senneman lakes
lhese hgures lo reecl realily, and re|ecls lhe ossibilily
of arlislic slylizalion, as lhe yramid shovs no signs
of mirror-image symmelry or 'olher kind of arls-con-
necled conslrainl' (. 165). Il is ossible, hovever, lhal
such a conslrainl exisled in lhe lolem oles sludied
by Marrion & Rosenblood (1986). They examined
11O deiclions of hand use in carved slanding oles
(nineleenlh lo lvenlielh cenlury) used for houses and
lolems in lhe Kvakiull Indian areas off lhe vesl coasl
of rilish Columbia. They found 2O er cenl righl-
handedness, 24 er cenl lefl-handedness, and 56 er
cenl simullaneous use of bolh hands.
|ngrctings
D'Lrrico (1992, 1OO, 99) reorls on four ossible vays
of conhguring lhe lvo hands and arms, derived from
his exerimenlal relicalions of engraved Azilian eb-
bles al Rochedane, le Mas d'Azil, and Iags, Irance.
The Azilian cullure vas Mesolilhic, belveen 11 kya
and 8 kya. D'Lrrico shoved (1988) lhal lhe engravings
vere made vorking lovards oneself, and lherefore
lhey lend lo roduce lhe frequenlly-observed allern
of righl lo lefl |uxlaosilion. bserving lhe clockvise
direclion of lurning vhile vorking lhe ebbles, he
noles lhal 'in all lhe engravings vilh lvo oosed
series lhe surface vas rolaled 18O degrees belveen
one series and lhe nexl' (1992, 1OO). This is a similar
conslrainl lo lhal menlioned by While (1998) for lhe
vorking of lvisled ovales. Sludying lhe grooves vilh
scanning eleclron microscoy, d'Lrrico (1988) shoved
lhal a righl-hander engraving from lefl lo righl creales
a groove vhich is comacled along lhe edge closesl
lo lhe user. Draving from lo lo bollom, lhe comacl
area runs along lhe lefl side (1988, 1726). The incisions
on lhe 27 Azilian ebbles are consislenl vilh having
been made from lefl lo righl by a righl-hander.
Prc|c !rcuing
There have been many suggeslions lhal, vhen drav-
ing a erson or animal in rohle, a righl-hander lends
lo drav lhe face lo lhe lefl, and a lefl-hander facing
righl. Iroonenls of lhis hyolhesis include Wilson
(1885, 132, 1891, 33), reuil (1952), Leroi-Gourhan
(1965), Ierell (197O, 141), Aller (1989), and Willcox
(1991, 146). ne of lhe earliesl menlions is from }.S.
187O (ciled in Uhrbrock 1973, 28), vho vrole
Mosl boys knov lhal il is easier lo drav a rohle vilh
lhe face looking lovard lhe lefl hand, yel on looking
over lhe hieroglyhs in lhe rilish Museum lhe faces
vill be generally found lovard lhe righl.
Aller (1989) made an exerimenlal sludy of 231
sub|ecls (19 L handers and 148 R) vho vere asked lo
quickly drav six shaes (bicycle, dog, bus, face rohle,
airlane, ilcher). The resulls shoved consislenl }-dis-
lribulions in lhe localion of lhe leading fealure (i.e. lhe
'direclion' of lhe draving). Namely, lhe direclion vas
slrongly correlaled vilh handedness, vilh mosl sub-
|ecls consislenlly direclional and a shar dro-off lo
veaker and veaker consislency. Like Wilson, Ierell
(197O) conhrms lhis lendency lo drav leading fealures
on lhe side conlralaleral lo lhe draving hand:
We have observed lhal vhen righl-handed children
drav man or animal faces, lhese faces alvays look
lefl. n lhe conlrary, lefl-handed children drav faces
looking righl. (Ierello 197O, 141)
The only one so far lo argue for more lefl-facing (and
hence more righl-handed arlisls), Wilson (1886, 17)
refers lo hieroglyhs al Ialenque (Mexico, Mayan
sile), in vhich 'mosl' of lhe animals are deicled look-
ing lo lhe lefl.
n lhe oosile side, some argue lhal rehisloric
ainlers vere more lefl-handed lhan novadays, or
ambidexlrous. This is lhe case of Ierell, reuil, Leroi-
Gourhan, and Willcox. reuil (1952) reorled hnding
5O.56 er cenl righl-orienled rohles in Luroean cave
arl (n 72O) (ciled in Willcox 1991). Similarly, Leroi-
Gourhan (1965) reorled 58.9 er cenl righl-orienled
rohles in Luroean cave arl (ciled in Willcox 1991).
Ierell found, in lhe Sanish cave of Allamira, equal
numbers of righl- and lefl-facing bison (Ierello 197O,
142). Irom lhese and olher references (books on cave
arl, elc.) Ierell agrees vilh lhe high roorlions of
lefl-handers among rehisloric 'arlisls'. Andr Leroi-
Gourhan (1965, ciled in Willcox) found 58.9 er cenl
righl-facing animals, as did reuil (1952), vho found
5O.56 er cenl righl-facing rohles (reuil (1952). Irom
a survey of several sile records, Willcox (1991) found
hgures ranging from 53 er cenl lo 64 er cenl righl-
facing animals in Soulh African, San, and nearby rock
arl. He seculales lhal lhe righl hemishere's sueri-
orily in face1allern recognilion, colour ercelion,
and olher visual abililies causes a higher roorlion
of lefl-handers lo be arlisls, and agrees lhal lhe Afri-
can dala suorls grealer rehisloric lefl-handedness
lhan in living San bushmen. Iales, ciled in Delluc &
232
Claplor 15
Delluc (1993, 44) also found more animals facing lo
lhe righl, and concluded lhere musl have been more
lefl-handers in rehislory.
Hcn!prinis cn! ncn! sicnci|s
A subslanlial body of lileralure has been ublished
on counls of righl and lefl handrinls and slencils in
caves. As lhese are frequenl lhroughoul lhe vorld,
easy lo counl, and vere direclly crealed by lhe hands
of aclual rehisloric eole, lhey conlribule valuable
dala. There are lhree vays of making reresenlalions
of hands in caves or rock shellers: 1) by diing lhe
hand in ainl, or ainling a molif onlo lhe hand, lhen
ressing lhe hand againsl lhe vall, 2) by ulling lhe
hand on lhe vall and sraying, or oullining, or dab-
bing ainl around il, 3) by draving a hand, eilher
slylized or realislic-looking. The hrsl melhod resulls
in osilive handrinls, lhe second melhod roduces
negalive hand slencils, and lhe lhird vill nol be con-
sidered here because il is disconnecled from any real,
aclual human hand.
Handedness can be inlerreled lhrough lhese
images if ve make lhe basic assumlion lhal lhe non-
dominanl hand vas referenlially laced on lhe vall,
and lhal il vas laced alm dovn. If lhe dominanl
hand vas needed lo hold lhe ainl alelle, lhe bloving
lube, lhe candle or lorch (since mosl cave ainlings
vere made beyond lhe lighl of day), or anylhing else,
lhen lhe nondominanl hand vould nalurally have
been lhe one selecled lo ress againsl lhe vall. When
using a bloving lube lo sray igmenl, lhe imlied
biomechanical conslrainl is lhal lhe dominanl hand
(lhe one vilh more recise conlrol) vill hold lhe lube,
leaving lhe nondominanl hand free lo be ainled on.
Wilhoul a lube, one mixes igmenl ovder vilh valer
and1or saliva and srays il direclly from lhe moulh.
Il may be necessary lo hold a conlainer of igmenl,
for vhich lhe dominanl hand mighl be used. Lven
vilhoul holding anylhing in lhe olher hand, il aears
more nalural lo ress lhe nondominanl hand againsl
lhe vall (see belov Gilaberl ers. comm., Iaurie &
Raymond 2OO4).
Ior osilive rinls, lhere is no queslion lhal lhe
alm musl have been dovn, since lhe skin on lhe back
of lhe hand has ils ovn dislinclive allern vhich is
nol lhal of a alm rinl. Ior negalive images or slen-
cils, in vhich ve only see lhe oulline of lhe hand, il
is ossible lo dislinguish belveen alm u and alm
dovn by several melhods: lhe clarily of lhe oulline,
lhe sreading angle belveen hngers, lhe resence
of lhe forearm, and lhe osilion of lhe hands on lhe
cave vall. Gradn (1994, 153) exlains lhal a dislincl
conlour can only be achieved vilh lhe alm dovn, lhe
back of lhe hand cannol be alied vilh enough res-
sure on lhe vall lo make a lighl seal. In discussing lhe
raclice of making hngers look deliberalely dislorled
(such as missing hngers), Walsh (1983, 4) indicales lhal
some Auslralian Aborigines avoided lhe undersray
roblem 'eilher by greasing lhe hand lo make il hl
close lo lhe surface, or by lacing lhe back of lhe hand
againsl lhe rock and lhen holding lhe hngers dovn'.
arrire & Sueres (1993, 52) conducled exerimenls
shoving lhal a dorsal hand osilion, even vilh lhe
hngers held dovn, roduces lhis invasive igmenl in
belveen lhe hngers due lo lhe lack of a lighl seal, and
furlhermore lhal lhe hand cannol reach ils maximum
sread of hngers vhen il is dorsally laced. This is
one vay lo rule oul lhe ossibilily of faking missing
hngers: lhe maximal sread belveen hngers cannol
be faked if lhe hngers are benl. Kirchner (1959), from
her ovn exerimenls, concluded il vas ossible lo
relicale, vilh clear oullines, lhe Gargas hands (vilh
missing hngers) by bending lhe hngers.
In addilion, lhe very osilion and angle of
handrinls can rule oul lhe use of one hand. Iirsl, lhe
heighl above ground (usually eye-level: Henneberg &
Malhers 1994) conslrains lhe angle al vhich lhe elbov
can bend. Second, if arl of lhe oulline of lhe forearm
is resenl, lhen il is clear vhich hand vas used, bul
even vilhoul lhe forearm, some lefl hand slencils vilh
a benl vrisl coming from lhe lefl side (see for examle
lhe frieze of 21 aligned hands al Gua Ham, orneo:
Iage & Chazine 2OO1) can only have been made vilh
lhe lefl hand alm dovn. This kind of vrisl-bending
can be seen in many holograhs of hand slencils
around lhe vorld, and seems lo be a nalural effecl of
lrying lo make a verlical hand slencil (because lhe dis-
lance required lo blov igmenl from one's ovn moulh
is around 4O cm (arrire & Sueres 1993, 49) lhe elbov
musl bend oulvard from lhe body). Iurlhermore, il
aears more nalural for a righl-hander lo lace lhe
lefl hand againsl lhe vall. Il has been observed by
one righl-handed ainler vilh four years' exerience
making hand slencils, lhal il feels more comforlable
and nalural lo ress lhe lefl hand againsl a vall (C.
Gilaberl ers. comm.). This is because lhe body musl
lvisl slighlly lo osilion lhe lefl hand on lhe vall
vhile leaving lhe correcl dislance belveen moulh and
vall. This seems lo imly some sorl of biomechanical
reference on lorso lvisling.
arrire & Sueres (1993, 5O) give lvo examles
from Gargas Cave (Irance, Gravellian era, 2621 kya)
lhal vould have been imossible lo make vilh lhe
righl alm u. ne slencil is a hand laced horizon-
lally vilh lhe vrisl lo lhe lefl, localed 2 m above lhe
ground in a narrov niche. In order for lhe righl hand
lo be used alm u, lhe erson vould have had lo
hang uside-dovn vhile a second erson srayed lhe
233
Humans, Tools and Handedness
igmenl, and yel lhe niche vas only big enough for
one erson. The second examle is a lefl hand vilh
lhe arm exlending lo lhe elbov, orienled 45 degrees
lo lhe righl, localed benealh a sloing vall.
The lechniques and melhods for making nega-
live and osilive hand images have been exlensively
sludied exerimenlally (e.g. Ringol 2OO2). The lech-
nique used 3O,OOO years ago can be delermined by
microscoic fealures of lhe igmenl on lhe cave vall,
such as degree of igmenl invasion inlo lhe orous
surface, lhe size of igmenl arlicles, and lheir dislri-
bulion (Clol ci c|. 1995). These aulhors idenlihed lhree
lechniques used al Gargas, for differenl colours of
igmenl. Ior red and yellov, and black carbon-based
igmenl slencils, liquid ainl vas alied by sray-
ing from lhe moulh. Ior black manganese slencils,
lhe igmenl vas alied by dabbing vilh a brush or
iece of fur (Clol ci c|. 1995, 231).
Iaunero (1992, 53), in a sludy aiming lo relicale
lhe negalive hands (or slencils) al Caadn de Los
Toldos, Ialagonia, exerimenlally lesled numerous
variables in order lo creale lhe negalive hand ainling
rocess vhich gives resulls 'mosl closely aroximal-
ing lhe observed archaeological realily'. He lesled lhe
lechnique in vhich ainl is direclly srayed from lhe
moulh. The variables vere: ainl densily, velocily
of alicalion, quanlily of air, dislance belveen lhe
moulh and hand, lemeralure of lhe ainl, basal area
(area covered by srayed ainl), angle of disersion
(indicales lhe amlilude of sraying), angle of moulh
osilion (measured from O degrees for lhe horizonlal
lo lhe direclion of sraying). He concluded lhal lhe
Ialagonian hand slencils al Caadn de Los Toldos
vere made vilh moulh sraying.
The melhod of sraying vilh a lube vas exeri-
menlally lesled by Iaurie & Raymond (2OO4). They
lesled 179 naive sub|ecls for lhroving hand, and
inslrucled lhem lo make a negalive hand using a en
vhich srays ink vhen blovn inlo, mimicking lhe
lechnique of rehisloric ainling vilh a lube. Iinally
lhe sub|ecls vere asked vhich vas lheir vriling hand.
Allhough mosl sub|ecls held lhe lube in lhe same hand
as lhe lhroving and vriling hand making slencils
of lhe oosile hand, as execled a surrising
number of righl-handers made righl-hand slencils
(17 er cenl). In conlrasl, only one lefl-hander made
a lefl-hand slencil. These resulls suggesl lhal lhe
archaeological record should shov more righl-hand
slencils lhan execled, if a cerlain roorlion of righl-
handers are adding lheir righl-hand oullines lo lhe
lolal of righl hand oullines roduced by lefl-hand-
ers. Iurlhermore, il can be deduced lhal lhe aclual
number of righl slencils indicales lhe very maximum
ossible number of lefl-handed eole, since some of
lhe images could have been roduced by righl-hand-
ers. Thus lhe number of lefl-handers can be eslimaled
from lhe number of righl slencils, if 93 er cenl of
righl slencils vere made by lefl-handers. Conversely,
nearly 1OO er cenl of lefl slencils can be allribuled lo
righl-handers. Comared vilh lhe summed dala from
Luroe, Iaurie & Raymond's roorlions are almosl
idenlical lo lhose of several Luroean caves, suggesl-
ing lhal similar roorlions of righl- and lefl-handers
exisled in lhe lasl 1O kya. Aside from lhe assumlion
lhal ainlers only ainled lheir ovn hands, Iaurie &
Raymond are also based on lhe suosilion lhal nega-
live hands vere ainled vilh a bloving lube. (arrire
& Sueres 1993, 5O describe a single ainling evenl in
Gargas, lvo adull hands framing a air of child's
hands, vhich imlies lhey vere ainled hands-free,
unless a lhird erson held lhe lube. The evidence for
sraying direclly from lhe moulh suggesls lhal lhe use
of a lube musl nol be laken for granled).
Taking inlo accounl lhe assumlions lhal lhe re-
ferred vay of making hand images vas alm-dovn
lacing of lhe nondominanl hand, ve can nov lurn lo
lhe archaeological dala. The oldesl cave ainlings are
currenlly in Chauvel Cave (Irance), and are daled lo
323O kya (Valladas ci c|. 2OO1, Clolles n.d. vebsile).
In lhe Cosquer cave (Irance, daled lo 2719 kya:
Valladas ci c|. 2OO1), Clolles & Courlin (n.d. vebsile)
counled 54 negalive hand slencils and one osilive
rinl. The roorlion of lefl and righl hands is un-
clear, bul lhere is a anel vilh 8 lefl slencils srayed
in black.
Delluc & Delluc (1993, 345) reviev lhe lileralure
from Irance and Sain, hnding ma|orilies of lefl hands
everyvhere: 17 caves vilh 319 hands yield 228 lefl
hands for 52 righl hands. In Kirchner's sludy (1959,
11O) on hand reresenlalions in lhe Iranco-Canlabrian
area, combined vilh dala from lhe resl of rehisloric
Luroe and elhnograhic dala from Auslralia and lhe
Americas, she counled 3O4 lefl-hand slencils for 71
righl, and 83 righl-hand rinls for 15 lefl-hand rinls.
This makes a lolal of 473 hands, consisling of 82 er
cenl righl-handedness and 18 er cenl lefl-handed-
ness. Kuhn (1955) had already counled 9 righl- and 35
lefl-hand slencils al lhe sile of Ll Caslillo, in Sain.
In a sludy of several caves on lhe Nullarbor Ilain,
Auslralia, Lane & Richards (1966, 46) reorl lhal 'lhe
ma|orily' of slencils are lefl hands. The slencils are
found in Murravi|inie Cave Numbers 1 and 3, Knov-
les Cave, and Abrakurrie Cave. The dales given for
nearby Koonalda Cave are 13,7OO and 18,2OO ya.
Iage and colleagues (Iage & Chazine 2OO1, Iage
ci c|. 2OO2) reorl on recenl discoveries in Kalimanlan,
orneo. There are 15OO hands in 26 caves, vhich have
been daled lo aboul 99OO ya (Ilagnes ci c|. 2OO3). The
234
Claplor 15
samle laken from a calcile draery covering a hand
slencil al Gua Saleh vas daled lo 987O ya, so lhe
image musl redale lhe draery. Tolal hand slencil
counls for one of lhe caves, Gua Tevel level 1, are
189. f lhese, 114 are lefl hands, 5O righl hands, and
25 indelerminale. This follovs lhe general allern of
aboul lvo-lhirds lefl hands lo one-lhird righl hands
in all lhe Kalimanlan caves (L.-H. Iage ers. comm.).
The magnihcenl frieze al Gua Ham, for examle, is
made of 21 aligned hands, of vhich 2O are lefl and 1
righl. The only excelion is in one seclion of lhe 'Tree
of Life' in Gua Tevel, consisling of 2O righl and 2 lefl
hands. The uniqueness of orneo's hand images are
lhe comosilion of inlerconnecled 'lrees' of hands,
connecled vilh lines, and decoraled vilh symbols
ainled onlo lhe alms.
Gradn (1994, 153) sludied lhe Ro Iinluras
(Iainled River) region of cave arl in Cenlral Ialago-
nia. Hand slencils dale lo al leasl 9 kya and exlend lo
53OO ya. Tvo caves, Cueva de Las Manos and Cueva
Grande of lhe Arroyo Ieo, yielded counls of 329 lefl
hands lo 31 righl ones, and 97 lefl hands for 2 righl
hands, reseclively.
Greer & Greer (1999, 6O) reorl on 7O8 rock-arl
siles in Monlana, USA. These caves, bluff faces and
rock shellers can be daled as beginning in lhe Middle
Archaic eriod (3OOO ) and exlending lo 14OO.
The ma|orily are osilive rinls (only 7 hand slencils
exisl). There are 429 idenlihable rinls vilh resecl lo
laleralily, 317 (74 er cenl) are righl hands and 112 (26
er cenl) are lefl hands.
Gunn (1998) describes lvo rock-arl comlexes in
lhe Levy Ranges, Auslralia, Kuli Mara and Irlikiri.
They have unique allerned handrinls vhich vere
made by hrsl ainling allerns onlo one hand, lhen
ressing lhe ainled hand onlo lhe rock surface. In 7
siles lhere are 1O9 allerned handrinls, 84 hand slen-
cils, and 7 handrinls. Reliable counls reveal 4:1 righl
lo lefl rinls, 1:4 righl lo lefl slencils, and generally
more lefl- lhan righl-allerned rinls. Gunn rooses
lhis laller facl reveals lhe lendency for righl-handers
lo ainl vilh lhe righl hand onlo lhe lefl. Suggesled
dales for lhese siles are fairly recenl, i.e. vilhin lhe
lasl 2OOO years, allhough one sheller vas occuied in
lhe lasl 3O kya.
Cnnc!usInns
We shall conclude by discussing lhe queslion lhal
slruclured lhis symosium namely, is knaing
slone a uniquely hominin behaviour` Il seems lhal
humans roulinely use a single 'referred' hand lo lay
lhe leading role in lool use, and lhal nearly nine limes
oul of len lhe choice is of lhe righl hand. As ve have
menlioned, and as is discussed by olher conlribulors
lo lhis volume, lhere is only veak and inconclusive
evidence for a similar oulalion-level bias lovards
lhe righl hand in lhe lool-using behaviours of olher
living rimales. This suggesls lhal lhere may also
be somelhing unique aboul lhe organizalion of lhe
neural subslrale lhal conlrols lhese volunlary manual
aclions. If ve analyze lhe skelelal and malerial cul-
lural evidence for a oulalion-level bias lovards
righl-handedness, ve see lhal lhe bias is consislenlly
observed in lhe remains of analomically modern hu-
man oulalions, and lhal lhere is a limiled quanlily
of evidence vhich consislenlly indicales lhal such a
bias also exisled among lhe earlier members of our
ovn genus. Whelher lhis reecls seleclion for lool-
making and lool-using caacilies, or seleclion for some
olher adalive caacily, is unforlunalely beyond lhe
scoe of lhis aer.
Allhough lhe sludy of human handedness cer-
lainly oens an inlriguing vindov onlo lhe asl, and
onlo lhe evolulion of human lool making and lool use,
lhere are olher queslions aboul hand skill vhich also
need lo be asked. Il is logically quile ossible lhal lhe
evolulion of handedness addresses only a arl of lhe
roblem of lhe evolulion of skill and comlexily in
lhe cncinc cprcicirc. If lhe evolulion of a bias lovards
righl-handedness receded lhe evolulion of a fully-
modern caacily for comlex serial order in lanning
and execuling lool-making and lool-using aclions,
lhen ve may be over-eslimaling lhe signihcance of
handedness as an evolulionary marker of 'lefl hemi-
shere' execulive funclions of a fully human kind,
including linguislic abilily. An inlegraled assessmenl
of bolh dimensions of lhe organizalion of skilled lool-
using aclions in lhe archaeological record is lhus long
overdue. Ior some handedness researchers lhis could
amounl lo 'lhinking lhe unlhinkable', bul lhinking
lhe unlhinkable is alvays a slimulaling (if unsellling)
exercise.
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