A Comprehensive Analysis of Long Bone Curvature in Neanderthals and Modern Humans Using 3D morphometrics Isabelle Elisabeth Peter Maria De Groote

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1 A Compehensive Analysis of Long Bone Cuvatue in Neandethals and Moden Humans Using 3D mophometics Isabelle Elisabeth ete Maia De Goote A thesis submitted to Univesity College London in fulfillment of the equiements fo the degee of Docto of hilosophy. 2008

2 I, Isabelle De Goote, confim that the wok pesented in this thesis is my own. Whee infomation has been deived fom othe souces, I confim that this has been indicated in the thesis. ii

3 Abstact Since thei discovey Neandethals wee descibed as having a maked degee of anteoposteio cuvatue of the femoal shaft. Although initially believed to be pathological, subsequent discoveies of Neandethal emains made femoal cuvatue as well as the lateal cuvatue of the adius to be consideed deived Neandethal featues. Femoal cuvatue has peviously been used in acial identification in moden humans but its functional significance is pooly undestood. A ecent study on Neandethals and ealy moden humans found no diffeences in femoal cuvatue, but did not conside size-coected cuvatue. Theefoe, the objectives of this study wee to 1) use 3D mophometic landmak and semi-landmak analysis to quantify bone cuvatue (femu, ulna, adius) in Neandethals, Uppe alaeolithic and ecent moden humans, 2) compae adult bone cuvatue between these populations, and 3) test hypotheses on the effects of climate, body size, and activity pattens on cuvatue. Compaisons between and within populations wee made using geometic mophometics (3D landmaks) and standad multivaiate methods. Compaative mateial involved all available Neandethal and Uppe alaeolithic moden human femoa, ulnae and adii, achaeological (Mesolithic, Neolithic, Medieval) and ecent human populations epesenting a wide geogaphical and lifestyle ange. The study found that thee ae significant diffeences in the anatomy of the femu, ulna and adius between Neandethals and moden humans. Neandethals have moe cuved femoa and adii than moden humans. Ealy moden humans ae most simila to ecent moden humans in thei anatomy. Recent moden human analyses indicate that femoal cuvatue and foeam cuvatue ae esponses to dispaate influences. Femoal cuvatue is a good indicato of activity level and habitual loading of the lowe limb. Cuvatue of the foeam is a consequence of cold adaptation and its pupose is to maintain biomechanical function of the foeam despite its foeshotening. iii

4 In memoy of Chalie. You knew my stengths, You knew my weaknesses, You wee my mento, You wee my fiend. iv

5 Acknowledgements Fist and foemost I thank my supevisos, D. Chales Lockwood and ofesso Leslie Aiello, who have been as much fiends as mentos. I also thank ofesso Simon Hillson fo his valuable advice. I am gateful to all the people who povided access to the fossils examined fo this thesis: I. Tattesal, G. Sawye at the Ameican Museum of Natual Histoy New Yok; R. Kuzynski at the Natual Histoy Museum, London; Y. Rak of the Sackle School of Medicine, Tel Aviv;. Mennecie at the Musée de l Homme, ais; J.-J. Cleyet-Mele at the National Museum of ehistoy, Les Eyzies; V. Melin-Anglade at the Museum of éigod, éiguex; M. Teschle- Nicola at the Natual Histoy Museum, Vienna;. Semal of Royal Belgian Institute of Natual Sciences, Bussels; W. Menghin at the Museum of ehistoy, Belin; J. Svoboda of the Institute of Achaeology, Dolni Vestonice; M. Dockalova at the Moavian Museum, Bno;. Velemínský, National Museum ague; J. Chistov at the Kunstkamea, Saint etesbug; D. ezhèmsky, Moscow State Univesity; M. Mogan at the eabody Museum, Havad Univesity, Boston; D. Hunt at the National Museum of Natual Histoy, Smithsonian Institute, Washington;. Bennike at the Medical Univesity, Copenhagen. I would like to thank hilip Mitteoecke and hilip Gunz fo the use of thei Mathematica functions and Katya Bulygina fo he patience in shaing he geometic mophometics knowledge. At Univesity College London (UCL) fist thanks go to the Anthopology Depatment Staff fo thei suppot thoughout my gaduate caee. I thank all my fiends ove the yeas at UCL who povided stimulating convesations, poof eadings, much needed distactions and emotional suppot. I thank all my non-ucl fiends and in paticula the Buddens, the Hubbads and the Madaas fo thei suppot and fo letting me be pat of thei families. Special thanks go to Ruth Bown who is the best English mum I could possibly have found. Finally, I thank my paents, Geetje and Alexande De Goote, fo suppoting me in the choices I make and fo encouaging me to fulfil my potential. I also thank my siste, Sabine, fo keeping me with both feet on the gound, and my bothe, Bet, fo his andom chats. I dedicate this thesis to my gandfathe, Edouad Van Hove, as without his wods of wisdom and funding I would not have been able to embak on this hd. This poject was suppoted by the Gaduate School, UCL; The Leakey Fund; The Ruggles-Gates Fund fo Biological Anthopological Reseach; The Wenne-Gen Foundation fo Anthopological Reseach; Synthesys; and the UCL Alumni. v

6 Table of contents CHATER 1. Intoduction upose of the study Long bone cuvatue Neandethals and moden humans Layout of the thesis... 9 CHATER 2. Histoy of eseach on long bone cuvatue Femu Compaative anatomy of the femu Intaspecific vaiation in femoal cuvatue Biomechanics acting on femoal cuvatue Radius and ulna Compaative anatomy of lowe am anatomy Intaspecific vaiation in the adius and ulna Biomechanics acting on lowe am cuvatue ossible causes fo vaiation in long bone cuvatue Neandethals and ickets Biomechanics and bone emodelling Body size Activity levels Climate Hypotheses and pedictions CHATER 3. Mateials and Methods Mateials Neandethal and ealy anatomically moden human fossils Neandethals Ealy moden humans Moden populations Methods opulation data and categoies Time peiod Envionmental data vi

7 Activity levels and subsistence stategy Individual data Univaiate measuements Bone shape Equipment and softwae Data acquisition and specimen set-up Landmaks and semi-landmaks Analytical methods Size adjustment fo the linea measuements ocustes methods Teatment of semi-landmaks incipal component analysis Inta-obseve eo Disciminant function analyses Analysis of Vaiance (ANOVA) Mantel test Othe univaiate analyses Ode of analysis fo Chapte Shape data Coelations between shaft shape and univaiate measuements Body size Sex Age Activity pattens Climate and latitude Evolution ove time Mantel test Systemic influences CHATER 4. Intaspecific diffeences in long bone cuvatue in moden humans Objective The femu Femu shape pincipal components explained Anteio suface (acuve) osteio suface (pcuve) vii

8 Medial suface (mcuve) Lateal suface (lcuve) oximal and distal epiphyses (Epi) Summay Coelations between Cs and univaiate measuements Factos influencing cuvatue in moden humans Body Size Sex Age Activity levels Evolution ove time in Euope Climate and latitude Mantel test Summay The lowe am Radius pincipal components explained Medial suface (mcuve) Lateal suface (lcuve) Epiphyses (Epi) Summay Ulna pincipal components explained osteio suface (pcuve) oximal ulna (pox) Summay Coelations between Cs and univaiate measuements Factos influencing cuvatue in moden humans Bilateal asymmety of the lowe am Body size Sex Age Activity levels Evolution ove time in Euope Climate and latitude Mantel test viii

9 Summay Systemic effects of cuvatue Discussion CHATER 5. Long bone cuvatue in Neandethals, Ealy and Recent moden humans Objective Femu Femu pincipal components explained Anteio suface (acuve) osteio suface (pcuve) Medial suface (mcuve) Lateal suface (lcuve) oximal and distal epiphyses (Epi) Summay Diffeences in femoal mophology between Neandethals, ealy and ecent moden humans Cuvatue Apex of cuvatue Othe shaft shape Epiphysis shape Univaiate measuements Disciminant function analysis Summay The lowe am Radius shape pincipal components explained Medial suface (mcuve) Lateal cuve (lcuve) Epiphyses (Epi) Summay The ulna pincipal components explained osteio cuve (pcuve) oximal ulna (pox) Summay Diffeences in lowe am mophology between Neandethals, ealy and ecent moden humans ix

10 Cuvatue Othe shaft shape Epiphyses shape Univaiate measuements Disciminant function analysis Summay CHATER 6. Discussion and Conclusion Discussion Conclusion REFERENCES AENDIX x

11 List of tables Table 3-1 Summay of the Neandethal sample, by egion Table 3-2 Summay of Ealy Moden Human sample, by egion Table 3-3 Summay of ecent moden human sample, alphabetically Table 3-4 Summay of individual data collected duing this poject Table 4-1 eason s coelation matix: femoal cuvatue Cs (n= 428) Table 4-2 eason s coelation matix femoal apex of cuvatue Cs (N=428) Table 4-3 eason s coelation matix fo femoal cuvatue Cs and univaiate measuements fo all moden human populations (N=36) Table 4-4 eason s coelation matix fo apex of cuvatue Cs and univaiate measuements fo all moden human populations (N=36) Table 4-5 eason s coelation matix fo othe shaft shape Cs and univaiate measuements fo all moden human populations (N=36) Table 4-6 eason s coelation matix fo femoal epiphyses shape Cs and univaiate measuements fo all human populations (N=36) Table 4-7 eason s coelation matix fo femoal cuvatue and univaiate measuements fo populations with high activity levels (N=21) Table 4-8 eason s coelation matix fo femoal apex of cuvatue and univaiate measuements fo populations with high activity levels (N=21) Table 4-9 eason s coelation matix fo othe femoal shaft shape Cs and univaiate measuements fo populations with high activity levels (N=21) Table 4-10 eason s coelation matix fo femoal epiphyses shape Cs and univaiate measuements fo populations with high activity levels (N=21) Table 4-11 eason s coelations fo body size (head diamete) and obusticity on the femu (N=36) Table 4-12 eason s coelations fo body size (head diamete) and obusticity on the femu (N=36) Table 4-13 Student s t-test esults fo obusticity in moden human males and females Table 4-14 Student s t-test esults fo femoal cuvatue in moden human males and females Table 4-15 Student s t-test esults fo obusticity in moden humans with high activity levels.109 Table 4-16 Student s t-test esults fo obusticity in moden humans with modeate activity levels xi

12 Table 4-17 Student s t-test esults fo obusticity in moden humans with low activity levels. 109 Table 4-18 Student s t-test esults fo cuvatue in moden humans with high activity levels Table 4-19 Student s t-test esults fo cuvatue in moden humans with modeate activity levels Table 4-20 Student s t-test esults fo cuvatue in moden humans with low activity levels Table 4-21 Student s t-test esults fo apex of cuvatue in moden human males and females.110 Table 4-22 Student s t-test esults fo apex of cuvatue in moden humans with high activity levels Table 4-23 Student s t-test esults fo apex of cuvatue in moden humans with modeate activity levels Table 4-24 Student s t-test esults fo apex of cuvatue in moden humans with low activity levels Table 4-25 Student s t-test esults fo othe aspects of shaft shape in moden human males and females Table 4-26 Student s t-test esults fo epiphysis shape in moden human males and females Table 4-27 Student s t-test esults fo epiphysis shape in moden humans with high activity levels Table 4-28 Student s t-test esults fo epiphysis shape in moden humans with modeate activity levels Table 4-29 Student s t-test esults fo epiphysis shape in moden humans with low activity levels Table 4-30 Student s t-test esults fo univaiate measuements in moden human males and females Table 4-31 Student s t-test esults fo univaiate measuements in moden humans with high activity levels Table 4-32 Student s t-test esults fo univaiate measuements in moden humans with modeate activity levels Table 4-33 Student s t-test esults fo univaiate measuements in moden humans with low activity levels Table 4-34 Kendall s Tau b coelations fo Cs and age (N=88) Table 4-35 Kendall s Tau b coelations fo univaiate measuements and age (N=88) Table 4-36 ANOVA esults fo adult age categoies on cuvatue Cs (N=4) Table 4-37 ANOVA esults fo activity levels and femoal cuvatue Cs xii

13 Table 4-38 ANOVA esults fo high activity subsistence categoies and femoal cuvatue Cs Table 4-39 ANOVA esults fo activity levels and the apex of femoal cuvatue Cs Table 4-40 ANOVA esults fo subsistence categoies and the apex of femoal cuvatue Cs Table 4-41 ANOVA esults fo activity levels and the othe femoal shaft shape Cs Table 4-42 ANOVA esults fo subsistence categoies and the othe femoal shaft shape Cs.127 Table 4-43 ANOVA esults fo activity level and the univaiate measuements of the femu Table 4-44 ANOVA esults fo subsistence categoies and the femoal univaiate measuements Table 4-45 ANOVA esults fo activity level categoies and the femoal epiphyses shape Cs Table 4-46 ANOVA esults fo subsistence categoies and the femoal epiphyses shape Cs. 139 Table 4-47 ANOVA esults fo time peiod and the femoal cuvatue Cs Table 4-48 ANOVA esults fo time peiod and the femoal cuvatue Cs Table 4-49 eason s coelations fo cuvatue, apex of cuvatue, diaphyseal shape and epiphyses shape Cs and latitude (climate) on the femu (N=35) Table 4-50 eason s coelations fo femoal univaiate measuements and latitude (climate) on the femu (N=35) Table 4-51 eason s coelations fo cuvatue, apex of cuvatue, diaphyseal shape and epiphyses shape Cs and latitude (climate) on the femu in high activity goups (N=17) Table 4-52 Results of the Mantel tests pefomed fo envionmental distance matices - femu Table 4-53 eason s coelation matix: adial cuvatue Cs (n= 360) Table 4-54 eason s coelation matix: adial cuvatue and epiphyses Cs (n= 349) Table 4-55 eason s coelation matix: posteio suface and poximal ulna Cs (n= 347) Table 4-56 eason s coelation matix fo adius shape Cs and univaiate measuements fo all moden human populations (N=35) Table 4-57 eason s coelation matix fo ulna shape Cs and univaiate measuements fo all moden human populations (N=35) Table 4-58 eason s coelation matix fo ulna shape Cs and univaiate measuements fo all moden human populations (N=35) Table 4-59 eason s coelation matix fo adius Cs and univaiate measuements fo populations with high activity levels (N=20) xiii

14 Table 4-60 eason s coelation matix fo ulna Cs and univaiate measuements fo populations with high activity levels (N=19) Table 4-61: eason s coelation matix fo ulna Cs and univaiate measuements fo populations with high activity levels (N=19) Table 4-62 Student s t-test esults fo bilateal asymmety in adius shape in moden humans.175 Table 4-63 Student s t-test esults fo bilateal asymmety in ulna shape in moden humans Table 4-64 Student s t-test esults fo univaiate measuements of the adius in moden humans Table 4-65 Student s t-test esults fo univaiate measuements of the ulna in moden humans Table 4-66 eason s coelations fo body size (head diamete) and adial cuvatue (N=27). 177 Table 4-67 eason s coelations fo body size (head diamete) and ulna shaft shape (N=27).177 Table 4-68 Student s t-test esults fo obusticity in moden human males and females Table 4-69 Student s t-test esults fo adius cuvatue Cs in moden human males and females Table 4-70 Student s t-test esults fo adius obusticity in moden human males and females with high activity levels Table 4-71 Student s t-test esults fo adius cuvatue Cs in moden human males and females with high activity levels Table 4-72 Student s t-test esults fo adius shaft shape Cs in moden human males and females ight only Table 4-73 Student s t-test esults fo adius shaft shape Cs in moden human males and females with high activity levels ight only Table 4-74 Student s t-test esults fo ulna shaft shape Cs in moden human males and females ight only Table 4-75 Student s t-test esults fo ulna shaft shape Cs in moden human males and females with high activity levels ight only Table 4-76 Student s t-test esults fo adius epiphyses Cs in moden human males and females ight only Table 4-77: Student s t-test esults fo adius epiphyses Cs in moden human males and females with high activity levels ight only Table 4-78 Student s t-test esults fo the poximal ulna Cs in moden human males and females ight only xiv

15 Table 4-79 Student s t-test esults fo the poximal ulna Cs in moden human males and females with high activity levels ight only Table 4-80: Student s t-test esults fo the univaiate measuements of the adius in moden human males and females. Undelined vaiables show bilateal asymmety and wee analysed fo the ight side only Table 4-81 Student s t-test esults fo the univaiate measuements of the adius in moden human males and females with high activity levels. Undelined vaiables show bilateal asymmety and wee analysed fo the ight side only Table 4-82 Student s t-test esults fo the univaiate measuements of the ulna in moden human males and females. Undelined vaiables show bilateal asymmety and wee analysed fo the ight side only Table 4-83 Student s t-test esults fo the univaiate measuements of the ulna in moden human males and females with high activity levels. Undelined vaiables show bilateal asymmety and wee analysed fo the ight side only Table 4-84 Kendall s Tau b coelations fo adius Cs and age (N=93) Table 4-85 Kendall s Tau b coelations fo ulna Cs (N=97) Table 4-86 Kendall s Tau b coelations fo the univaiate measuements on the adius and age (N=97) Table 4-87 Kendall s Tau b coelations fo univaiate measuements on the ulna (N=97) Table 4-88 ANOVA esults fo age categoies and adius cuvatue Cs Table 4-89 ANOVA esults fo age categoies and ulna shape Cs Table 4-90 ANOVA esults fo activity levels and adius cuvatue Cs Table 4-91 ANOVA esults fo high activity subsistence subsistence stategies and adius cuvatue Cs Table 4-92 ANOVA esults fo activity levels and adius shaft shape Cs Table 4-93 ANOVA esults fo subsistence stategies with high activity levels and adius shaft shape Cs Table 4-94 ANOVA esults fo activity levels and ulna shaft shape Cs Table 4-95 ANOVA esults fo subsistence stategies with high activity levels and ulna shaft shape Cs Table 4-96 ANOVA esults fo activity levels and adius epiphyses Cs Table 4-97 ANOVA esults fo subsistence goups with high activity levels and adius epiphyses Cs Table 4-98 ANOVA esults fo activity levels and the poximal ulna Cs xv

16 Table 4-99 ANOVA esults fo subsistence stategy and the poximal ulna Cs Table ANOVA esults fo activity level and adius obusticity Table ANOVA esults fo subsistence stategy and adius obusticity Table ANOVA esults fo activity level and univaiate measuements on the adius Table ANOVA esults fo activity level and univaiate measuements on the adius ight only Table ANOVA esults fo subsistence stategy and univaiate measuements on the adius Table ANOVA esults fo activity levels and univaiate measuements on the ulna Table ANOVA esults fo subsistence pattens and univaiate measuements on the ulna Table ANOVA esults fo time-peiod and cuvatue of the adius Table ANOVA esults fo time-peiod and ulna shape Table eason s coelations fo cuvatue, apex of cuvatue, diaphyseal shape and epiphyses shape Cs and latitude (climate) on the adius (N=34) Table eason s coelations fo cuvatue, apex of cuvatue, diaphyseal shape and epiphyses shape Cs and latitude (climate) on the ulna (N=32) Table eason s coelations fo adius obusticity (head, midshaft and distal aticulation) and latitude (climate) (N=34) Table eason s coelations fo univaiate measuements on the adius and latitude (climate) (N=34) Table eason s coelations fo univaiate measuements on the ulna and latitude (climate) (N=31) Table Results of the Mantel tests pefomed fo envionmental distance matices adius Table Results of the Mantel tests pefomed fo envionmental distance matices - ulna239 Table eason s coelations fo cuvatue and apex of cuvatue Cs between the femu, adius and ulna (N=218) Table 5-1 eason s coelation matix: femoal cuvatue Cs (n= 449). Neandethals, ealy and ecent moden humans Table 5-2 eason s coelation matix: femoal apex of cuvatue Cs (n= 449). Neandethals, ealy and ecent moden humans Table 5-3 ANOVA esults fo palaeogoup and femoal cuvatue Cs Table 5-4 ANOVA esults fo palaeogoup and femoal apex of cuvatue Cs xvi

17 Table 5-5 ANOVA esults fo palaeogoup and othe femoal shaft shape Cs Table 5-6 ANOVA esults fo palaeogoup and othe femoal shaft shape Cs Table 5-7 Desciptives fo Neandethals, ealy and ecent moden humans and the univaiate measuements of the femu Table 5-8 ANOVA esults fo palaeogoup and femoal univaiate measuements Table 5-9 Disciminant function coefficients - femu Table 5-10 eason s coelation matix fo adius cuvatue Cs (n= 391) Table 5-11 eason s coelation matix fo adius cuvatue and epiphyses Cs (n= 377) Table 5-12 eason s coelation matix fo adius cuvatue and othe shaft shape Cs (n= 391) Table 5-13 eason s coelation matix fo adius epiphyses and othe shaft shape Cs (n= 377) Table 5-14 eason s coelation matix: ulna Cs (n= 344) Table 5-15 ANOVA esults fo palaeogoup and adius cuvatue Cs Table 5-16 ANOVA esults fo palaeogoup and othe adius shaft shape Cs Table 5-17 ANOVA esults fo palaeogoup and ulna shaft shape Cs Table 5-18 ANOVA esults fo palaeogoup and adius epiphysis shape Cs Table 5-19 ANOVA esults fo palaeogoup and poximal ulna Cs Table 5-20 Desciptives of palaeogoup and the univaiate measuements of the adius Table 5-21 ANOVA esults fo palaeogoup and univaiate measuements of the adius Table 5-22 Desciptives of palaeogoup and the univaiate measuements of the adius Table 5-23 ANOVA esults fo palaeogoup and univaiate measuements of the ulna Table 5-24 Disciminant function coefficients adius Table 5-25 Disciminant function coefficients - ulna xvii

18 List of figues Figue 2-1 Subtense method employed by Walensky (1962) Figue 2-2 An Afican-Ameican, Inuit and a Native Ameican femu (fom Walensky, 1965) showing inceasing amounts of cuvatue and loweing apexes of cuvatue Figue 2-3 Tudell s method of measuing cuvatue by placing the femu on two blocks (Tudell, 1999) Figue 2-4 Muscles acting on the femu Figue 2-5 Hominoid adii Figue 2-6 X-ay image of an infant with sevee ickets Figue 3-1 Specimen set up fo the femu, adius and ulna using clamps and a test tube stand.. 71 Figue 4-1 The fist and second Cs fo the anteio cuve of the femu. All ecent moden human samples Figue 4-2 Mophological tends fo the anteio cuve of the femu fo all ecent moden humans Figue 4-3 The fist and thid Cs fo the posteio cuve of the femu. All ecent moden human samples. Cs ae explained in Figue Figue 4-4 Mophological tends fo the posteio cuve of the femu fo all ecent moden humans Figue 4-5 The fist and second Cs fo the medial cuve of the femu. All ecent moden human samples. Cs ae explained in Figue 4-6Figue Figue 4-6 Mophological tends fo the medial cuve of the femu fo all ecent moden humans. All in lateal view Figue 4-7 The fist and second Cs fo the lateal cuve of the femu. All ecent moden human samples. Cs ae explained in Figue Figue 4-8 Mophological tends fo the lateal cuve of the femu fo all ecent moden humans Figue 4-9 Mophological tends fo the epiphyses of the femu fo all ecent moden humans. All anteio view Figue 4-10 Distibution of the activity level categoies in the space of C1 (degee of cuvatue) and C2 (apex of cuvatue) of the anteio cuve fo all moden humans Figue 4-11 Distibution of the activity level categoies in the space of C1 (degee of cuvatue) and C2 (apex of cuvatue) of the posteio cuve fo all moden humans xviii

19 Figue 4-12 Anteio femoal cuvatue fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-13 osteio femoal cuvatue fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-14 Anteio femoal apex of cuvatue fo moden humans, by activity level. Scale is evesed so that highe values indicate a moe poximal apex of cuvatue. Mean and 95% confidence inteval (whiskes) Figue 4-15 osteio femoal apex of cuvatue fo moden humans, by activity level. Scale is evesed so that highe values indicate a moe poximal apex of cuvatue. Mean and 95% confidence inteval (whiskes) Figue 4-16 Anteio femoal apex of cuvatue fo moden humans, by subsistence stategy. Scale is evesed so that highe values indicate a moe poximal apex of cuvatue. Mean and 95% confidence inteval (whiskes) Figue 4-17 cuvamhc4 fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-18 McuvAMHC3 fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-19 LcuvAMHC2 fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-20 LcuvAMHC4 fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-21 cuvamhc2 fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-22 Femu length fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-23 Neck-shaft angle fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-24 Subtochanteic shape atio fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-25 Midshaft shape atio fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-26 Subpilastic shape atio fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) xix

20 Figue 4-27 Neck-length atio fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-28 Robusticity index fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-29 Femoal head obusticity fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-30 Femu length fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-31 Neck-shaft angle fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-32 Tosion angle fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-33 Midshaft shape fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-34 Robusticity index fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-35 Femoal head obusticity fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-36 EpiAMHC2 (epiphysis width) fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-37 EpiAMHC5 (neck length) fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-38 EpiAMHC1 (epiphysis width) fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-39 EpiAMHC3 (tosion and distal epiphysis width) fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-40 EpiAMHC5 (neck length) fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-41 Distibution of time peiods in the space of C1 (degee of cuvatue) and C2 (apex of cuvatue) of the anteio cuve fo all moden humans Figue 4-42 Distibution of the time peiods in the space of C1 (degee of cuvatue) and C3 (apex of cuvatue) of the posteio cuve fo all moden humans Figue 4-43 Anteio femoal cuvatue fo moden Euopeans, by time peiod. Mean and 95% confidence inteval (whiskes) xx

21 Figue 4-44 osteio femoal cuvatue fo moden Euopeans, by time peiod. Mean and 95% confidence inteval (whiskes) Figue 4-45 Anteio apex of femoal cuvatue fo moden Euopeans, by time peiod. Mean and 95% confidence inteval (whiskes) Figue 4-46 Femu length and absolute latitude fo the ecent moden human sample including the small bodied equatoial samples: ygmy, euvian and Andamanese samples Figue 4-47 The fist and second Cs fo the medial cuve of the adius. All ecent moden human samples. Cs ae explained in Figue Figue 4-48 Mophological tends fo the medial cuve of the adius fo all ecent moden humans Figue 4-49 The fist and second Cs fo the lateal cuve of the adius. All ecent moden human samples. Cs ae explained in Figue Figue 4-50 Mophological tends fo the lateal cuve of the adius fo all ecent moden humans Figue 4-51 Mophological tends fo the epiphyses of the adius fo all ecent moden humans. All medial view Figue 4-52 The fist and second Cs fo the posteio cuve of the ulna. All ecent moden human samples. Cs ae explained in Figue Figue 4-53 Mophological tends fo the posteio cuvatue of the ulna fo all ecent moden humans Figue 4-54 The fist and second Cs fo the poximal ulna. All ecent moden human samples. Cs ae explained in figue Figue Figue 4-55 Mophological tends fo the poximal ulna fo all ecent moden humans Figue 4-56 oxamhc2 fo moden humans, by age stategy. Mean and 95% confidence inteval (whiskes) Figue 4-57 Distibution of the activity level categoies in the space of C1 (degee of cuvatue) and C2 (medial expansion of the inteosseous cest) of the medial cuve of the adius fo all moden humans. Cicles: high activity, squaes: modeate activity, cosses: low activity Figue 4-58 Distibution of the activity level categoies in the space of C1 (degee of cuvatue) and C2 (apex of cuvatue) of the lateal cuve of the adius fo all moden humans Figue 4-59 Distibution of the activity level categoies in the space of the C1 (degee of mediolateal cuvatue) and C2 (mediolateal sinusoidal shape) of the posteio cuve of the ulna fo all moden humans Figue 4-60 Lateal cuvatue of the adius fo moden humans, by subsistence stategy xxi

22 Figue 4-61 LcuvAMHC3 (high values have the least sinusoidal shaft) of the adius fo moden humans, by high activity subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-62 McuveC2 of the adius fo moden humans, by high activity subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-63 LcuveC3 (low values ae moe sinusoidal) of the adius fo moden humans, by high activity subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-64 EpiAMHC2 fo moden humans, by subsistence stategy. The scale of lcuveamhc1 is evesed to ease intepetations (have a moe posteioly oiented head). Mean and 95% confidence inteval (whiskes) Figue 4-65 oxamhc4 fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-66 oxamhc2 fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-67 oxamhc4 fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-68 Head obusticity fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-69 Midshaft obusticity fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-70 Head obusticity fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-71 Relative distal aticulation size fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-72 osition of the adial tubeosity fo moden humans, by activity level. Lowe values ae moe medially placed. Mean and 95% confidence inteval (whiskes) Figue 4-73 Neck-shaft angle fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-74 Relative adial neck length fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-75 Dosal subtense fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-76 Maximum length fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) xxii

23 Figue 4-77 Neck-shaft angle fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-78 osition of the adial tubeosity fo moden humans, by subsistence stategy. Lowe values ae moe medially placed. Mean and 95% confidence inteval (whiskes) Figue 4-79 Dosal subtense fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-80 Neck length atio fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-81 Midshaft shape atio fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-82 Maximum length fo moden humans, by activity level Figue 4-83 Midshaft shape fo moden humans, by activity level Figue 4-84 Radial notch suface aea fo moden humans, by activity level Figue 4-85 Tochlea notch oientation fo moden humans, by activity level Figue 4-86 Olecanon oientation angle fo moden humans, by activity level Figue 4-87 Coonoid-olecanon atio fo moden humans, by activity level Figue 4-88 Bachial muscle attachment atio fo moden humans, by activity level. Highe values have a elatively lowe insetion. Mean and 95% confidence inteval (whiskes) Figue 4-89 Robusticity at 50% shaft level fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-90 Robusticity at 25% shaft level fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes) Figue 4-91 Ulna maximum length fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-92 osition of the bachial tubeosity fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-93 Olecanon-shaft size atio fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-94 Radial notch suface atio fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-95 Tochlea notch oientation fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes) Figue 4-96 Olecanon oientation angle fo moden humans, by subsistence stategy Figue 4-97 Coonoid-olecanon atio fo moden humans, by subsistence patten xxiii

24 Figue 4-98 Relative size of the ponato cest fo moden humans, by subsistence patten Figue 4-99 Robusticity at 50% shaft level fo moden humans, by subsistence patten. Mean and 95% confidence inteval (whiskes) Figue Robusticity at 25% shaft level fo moden humans, by subsistence patten. Mean and 95% confidence inteval (whiskes) Figue Robusticity of the distal aticulation fo moden humans, by subsistence patten. Mean and 95% confidence inteval (whiskes) Figue Lateal cuvatue of the adius fo moden Euopeans, by time peiod Figue cuveamhc3 (high values have a moe anteoposteioly sinusoisal shaft) of the adius fo moden Euopeans, by time peiod Figue oxamhc4 (high values have a deepe tochlea notch) of the adius fo moden Euopeans, by time peiod Figue Lateal cuvatue of the adius (lcuveamhc1) and latitude fo ecent moden humans Figue Medial expansion of the inteosseous cest (mcuveamhc2) and latitude fo ecent moden humans Figue Sinusoidal shape of the adius (mcuveamhc3) and latitude fo ecent moden humans Figue Distance between the 80% shaft level and the tip of the coonoid pocess (poxamhc2) and absolute latitude fo ecent moden humans Figue Oientation of the tochlea notch (poxamhc3) and absolute latitude fo ecent moden humans Figue Radius midshaft obusticity and latitude fo ecent moden humans Figue Radius distal aticulation obusticity and latitude fo ecent moden humans Figue Radius head obusticity and latitude fo ecent moden humans Figue Scatteplot fo olecanon shaft atio and latitude fo ecent moden humans Figue Radial notch suface aea and latitude fo ecent moden humans Figue Oientation of the tochlea notch and latitude fo ecent moden humans Figue Olecanon oientation angle and latitude fo ecent moden humans Figue osition of the bachial muscle insetion and latitude fo ecent moden humans Figue Distal aticulation obusticity of the ulna and latitude fo ecent moden humans xxiv

25 Figue 5-1 Mophological tends fo the anteio cuve of the femu fo Neandethals, ealy and ecent moden humans Figue 5-2 Mophological tends fo the posteio cuve of the femu fo Neandethals, ealy and ecent moden humans Figue 5-3 Mophological tends fo the medial cuve of the femu fo Neandethals, ealy and ecent moden humans. All lateal view Figue 5-4 Mophological tends fo the lateal cuve of the femu fo Neandethals, ealy and ecent moden humans Figue 5-5 Mophological tends fo the epiphyses of the femu fo Neandethals, ealy and ecent moden humans Figue 5-6 The anteio cuve of the femu fo Neandethals, ealy and ecent moden humans. (Line=mean, Box= 2 S.E., whiskes: 2 S.D.). The highe values fo Neandethalss indicate that they ae moe cuved than the moden humans Figue 5-7 The posteio cuve of the femu fo Neandethals, ealy and ecent moden humans. (Line=mean, Box= 2 S.E., whiskes: 2 S.D.). The lowe values fo Neandethal indicates that they ae moe cuved than the moden humans Figue 5-8 The anteio apex of cuvatue of the femu fo Neandethals, ealy and ecent moden humans. (Line=mean, Box= 2 S.E., whiskes: 2 S.D.). The highe value fo Neandethals indicates a lowe apex of cuvatue Figue 5-9 LcuAllC3 fo Neandethals, ealy and ecent moden humans Figue 5-10 EpiAllC1 fo Neandethals, ealy and ecent moden humans Figue 5-11 EpiAllC2 fo Neandethals, ealy and ecent moden humans Figue 5-12 EpiAllC3 fo Neandethals, ealy and ecent moden humans Figue 5-13 EpiAllC5 fo Neandethals, ealy and ecent moden humans Figue 5-14 Femu length fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-15 Neck-shaft angle fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-16 Tosion angle fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-17 Subtochanteic atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-18 Midshaft atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) xxv

26 Figue 5-19 Subpilastic atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-20 Condyle obusticity fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-21 Neck length atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-22 Robusticity index fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-23 Head obusticity fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-24 Disciminant Function 1 and 2 fo Neandethals, ealy and ecent moden humans Figue 5-25 Mophological tends fo the medial cuve of the adius fo Neandethals, ealy and ecent moden humans Figue 5-26 Mophological tends fo the lateal cuve of the adius fo Neandethals, ealy and ecent moden humans Figue 5-27 Mophological tends fo the epiphyses of the adius fo Neandethals, ealy and ecent moden humans. All medial view Figue 5-28 The fist Cs fo the medial and lateal cuve of the adius Figue 5-29 Mophological tends fo the posteio cuve of the ulna fo Neandethals, ealy and ecent moden humans Figue 5-30 Mophological tends fo the poximal ulna fo Neandethals, ealy and ecent moden humans Figue 5-31 The medial cuve of the adius fo Neandethals, ealy and ecent moden humans Figue 5-32 The lateal cuve of the adius fo Neandethals, ealy and ecent moden humans t Figue 5-33 LcuAllC2 fo Neandethals, ealy and ecent moden humans Figue 5-34 LcuAllC3 fo Neandethals, ealy and ecent moden humans Figue 5-35 cuallc1 fo Neandethals, ealy and ecent moden humans Figue 5-36 cuallc2 fo Neandethals, ealy and ecent moden humans Figue 5-37 oxallc2 fo Neandethals, ealy and ecent moden humans Figue 5-38 oxallc3 fo Neandethals, ealy and ecent moden humans xxvi

27 Figue 5-39 Maximum adius length fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-40 osition of the adial tubeosity fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-41 Dosal subtense fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-42 Relative adius neck length fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-43 Head shape atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-44 Midshaft shape atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-45 Maximum ulna length fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-46 Olecanon-shaft atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-47 Midshaft shape atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-48 Radial notch suface aea fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-49 Tochlea notch oientation fo Neandethals, ealy and ecent moden humans. 310 Figue 5-50 Olecanon oientation fo Neandethals, ealy and ecent moden humans Figue 5-51 Coonoid-olecanon atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-52 Bachial insetion atio fo Neandethals, ealy and ecent moden humans. A highe value means a elatively lowe insetion. Mean and 95% confidence inteval (whiskes) Figue 5-53 Ulna obusticity at 50% shaft level fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-54 Robusticity of the head of the ulna aea fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Figue 5-55 Disciminant Function 1 and 2 fo fo Neandethals, ealy and ecent moden humans xxvii

28 Figue 5-56 Disciminant Function 1 and 2 fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) xxviii

29 List of appendices Appendix 1 Landmaks and measuements fo the femu Appendix 2 Landmak diagam femu (Afte ) Appendix 3 Landmaks and measuements fo the adius Appendix 4 Landmak and measuement diagams adius (Afte and ) Appendix 5 Landmaks and measuements fo the ulna Appendix 6 Landmak and measuement diagams ulna (Afte and ) Appendix 7 Diagams fo measuements calculated fom the landmaks on the ulna (Afte and ) Appendix 8 Rainfall distance matix between populations (data fom Hijmans, et al., 2005) Appendix 9 Tempeatue distance matix between populations (data fom Hijmans, et al., 2005) Appendix 10 Altitude distance matix between populations (data fom Hijmans, et al., 2005) 376 Appendix 11 ost-hoc compaisons fo activity levels and femoal cuvatue Cs. Matix fo paiwise mean diffeences between categoies Appendix 12 ost-hoc compaisons fo high activity subsistence stategies and femoal cuvatue Cs. Matix fo paiwise mean diffeences between categoies Appendix 13 ost-hoc compaisons fo activity levels and apex of cuvatue Cs. Matix fo paiwise mean diffeences between categoies Appendix 14 ost-hoc compaisons fo high activity subsistence stategies and femoal apex of cuvatue Cs. Matix fo paiwise mean diffeences between categoies Appendix 15 ost-hoc compaisons fo activity levels and othe femoal shaft shape Cs. Matix fo paiwise mean diffeences between categoies Appendix 16 ost-hoc compaisons fo high activity subsistence stategies and othe femoal shaft shape Cs. Matix fo paiwise mean diffeences between categoies Appendix 17 ost-hoc compaisons fo activity levels and femoal univaiate measuements. Matix fo paiwise mean diffeences between categoies Appendix 18 ost-hoc compaisons fo high activity subsistence stategies and femoal univaiate measuements. Matix fo paiwise mean diffeences between categoies Appendix 19 ost-hoc compaisons fo activity levels and femoal epiphysis shape Cs. Matix fo paiwise mean diffeences between categoies xxix

30 Appendix 20 ost-hoc compaisons fo high activity subsistence stategies and femoal epiphysis shape Cs. Matix fo paiwise mean diffeences between categoies Appendix 21 ost-hoc compaisons fo time peiod and femoal apex of cuvatue Cs. Matix fo paiwise mean diffeences between categoies Appendix 22 ost-hoc compaisons fo high activity subsistence stategies and adius cuvatue Cs. Matix fo paiwise mean diffeences between categoies Appendix 23 ost-hoc compaisons fo activity levels stategies and adius shaft shape Cs. Matix fo paiwise mean diffeences between categoies Appendix 24 ost-hoc compaisons fo high activity subsistence stategies and adius shaft shape Cs. Matix fo paiwise mean diffeences between categoies Appendix 25 ost-hoc compaisons fo activity levels and adius epiphysis shape Cs. Matix fo paiwise mean diffeences between categoies Appendix 26 ost-hoc compaisons fo high activity subsistence stategies and adius epiphysis shape Cs. Matix fo paiwise mean diffeences between categoies Appendix 27 ost-hoc compaisons fo activity levels and poximal ulna Cs. Matix fo paiwise mean diffeences between categoies Appendix 28 ost-hoc compaisons fo high activity subsistence stategies and poximal ulna Cs. Matix fo paiwise mean diffeences between categoies Appendix 29 ost-hoc compaisons fo activity levels and adius obusticity. Matix fo paiwise mean diffeences between categoies Appendix 30 ost-hoc compaisons fo high activity subsistence stategies and adius obusticity. Matix fo paiwise mean diffeences between categoies Appendix 31 ost-hoc compaisons fo activity levels and adius univaiate measuements. Matix fo paiwise mean diffeences between categoies Appendix 32 ost-hoc compaisons fo activity levels and adius univaiate measuements ight only. Matix fo paiwise mean diffeences between categoies Appendix 33 ost-hoc compaisons fo high activity subsistence stategies and adius univaiate measuements. Matix fo paiwise mean diffeences between categoies Appendix 34 ost-hoc compaisons fo activity levels and ulna univaiate measuements. Matix fo paiwise mean diffeences between categoies Appendix 35 ost-hoc compaisons fo high activity subsistence stategies and ulna univaiate measuements. Matix fo paiwise mean diffeences between categoies Appendix 36 ost-hoc compaisons fo time peiod and adius cuvatue Cs. Matix fo paiwise mean diffeences between categoies xxx

31 Appendix 37 ost-hoc compaisons fo time peiod and ulna Cs. Matix fo paiwise mean diffeences between categoies Appendix 38 ost-hoc compaisons fo palaeogoup and femual cuvatue Cs. Matix fo paiwise mean diffeences between categoies Appendix 39 ost-hoc compaisons fo palaeogoup and femoal apex of cuvatue Cs. Matix fo paiwise mean diffeences between categoies Appendix 40 ost-hoc compaisons fo palaeogoup and othe femoal shaft shape Cs. Matix fo paiwise mean diffeences between categoies Appendix 41 ost-hoc compaisons fo palaeogoup and femoal epiphysis shape Cs. Matix fo paiwise mean diffeences between categoies Appendix 42: ost-hoc compaisons fo palaeogoup and femoal univaiate measuments. Matix fo paiwise mean diffeences between categoies Appendix 43 ost-hoc compaisons fo palaeogoup and adius cuvatue Cs. Matix fo paiwise mean diffeences between categoies Appendix 44 ost-hoc compaisons fo palaeogoup and othe adius shaft shape Cs. Matix fo paiwise mean diffeences between categoies Appendix 45 ost-hoc compaisons fo palaeogoup and ulna shaft shape Cs. Matix fo paiwise mean diffeences between categoies Appendix 46 ost-hoc compaisons fo palaeogoup and poximal ulna Cs. Matix fo paiwise mean diffeences between categoies Appendix 47 ost-hoc compaisons fo palaeogoup and adius univaiate measuements. Matix fo paiwise mean diffeences between categoies Appendix 48 ost-hoc compaisons fo palaeogoup and ulna univaiate measuements. Matix fo paiwise mean diffeences between categoies xxxi

32 CHATER 1. INTRODUCTION 1.1. upose of the study When in the 19 th Centuy the Feldhofe Neandethal emains wee discoveed, eseaches noted a maked degee of anteio cuvatue of the femoal shaft and ascibed it to pathology (Klaatsch, 1901; Boule, 1908; Tinkaus and Shipman, 1993). With the subsequent discoveies of othe Neandethal emains, femoal cuvatue was consideed to be a deived featue of Neandethals as wee the shotened and cuved ulna and adius (Klaatsch, 1901; Tinkaus and Shipman, 1993; Chuchill, 1998; Golovanova et al., 1999; Czanetzki, 2000; Weave, 2003; Yamanaka et al., 2005). Relatively little wok has been done to quantify diaphyseal cuvatue in Neandethals, but a ecent study analysed pattens of femoal cuvatue in Neandethals, ecent humans and Late leistocene ealy moden humans (Shackelfod and Tinkaus, 2002). Shackelfod and Tinkaus (2002) suggested that Neandethals wee indistinguishable fom Middle alaeolithic and ealy Uppe alaeolithic ealy moden humans in thei degee of absolute anteio cuvatue. Additionally, most of the individuals in these alaeolithic populations wee found to exhibit a moe distal apex of cuvatue (point of maximum cuvatue) compaed to moe ecent populations (Shackelfod and Tinkaus, 2002). They suggested that this could be coelated with measues of bone hypetophy o an oveall decease in lowe-limb obusticity duing the Middle to Uppe alaeolithic. The five egional goups fom which thei samples oiginated wee significantly diffeent in femoal cuvatue and Shackelfod and Tinkaus (2002) suggested that the oveall decease in femoal cuvatue in moden humans was due to a decease in longdistance mobility. Reseach fom foensic anthopology also suggests that significant diffeences exist in femoal cuvatue between moden human populations (Stewat, 1962; Walensky, 1962, 1965; Gilbet, 1975, 1976; Tudell, 1999). Initial studies demonstated the diagnostic value of femoal cuvatue in distinguishing between Native Ameican, Afican-Ameican and Caucasoid Ameican populations (Stewat, 1962; Walensky, 1962, 1965; Gilbet, 1975, 1976; Tudell, 1

33 1999). When the eseach was expanded by inceasing the numbe of populations, no elationship was found between femoal cuvatue, habitual behavioual pattens and latitudinal position of those populations (Stewat, 1962; Walensky, 1962, 1965; Gilbet, 1975, 1976; Tudell, 1999). Tudell (1999) efined the measuement techniques by taking measuements at thee points along the cuve and found an 87.12% aveage accuate ace detemination fo Afican-Ameicans and Caucasoids (see Chapte 2 fo moe details). The moe detailed chaacteisation of cuvatue possible with 3D mophometics has the potential to efine the diffeences between moden human goups. The Neandethal adius has also been descibed as being moe lateally cuved than that of humans and to fall beyond the highe limits of moden human vaiation (Fische, 1906; Botez, 1926 in atte, 1955; Vandemeesch and Tinkaus, 1995; Caeteo et al., 1999; Czanetzki, 2000). Fische (1906) descibed Neandethals to have a lage posteio subtense in the ulna but moe ecent wok has not investigated this. In the eseach pesented hee, I conside the diffeences and similaities in long bone cuvatue and position of the apex of cuvatue of the femu, ulna and adius. This study has thee main objectives: 1) to detemine the influence of climatic, body size and behavioual coelates on the obseved diffeences in bone cuvatue in Holocene moden humans, 2) to descibe diffeences in long bone cuvatue between Neandethals and moden humans, and 3) to detemine how the factos that influence moden human bone cuvatue can be applied to infom ou undestanding of Neandethals and ealy moden humans. The fist objective involves an analysis of pattens of cuvatue and anthopometic measuements of moden humans and thei elationship to population-specific infomation such as body size, activity level, time peiod and climate. This will be done in ode to identify the biomechanical and adaptive advantages of diffeent degees of cuvatue within moden humans, in ode to fom pedictions fo the degee of cuvatue obseved in Neandethals and ealy moden humans. The second objective equies an analysis to test whethe thee ae any significant diffeences between Neandethals and moden humans in femoal and lowe am cuvatue. The long claimed distinction in degee of femoal cuvatue in Neandethals was challenged by Shackelfod and Tinkaus (2002) who found no diffeence between Neandethals and moden 2

34 humans. This hypothesis will be tested again hee on the cuvatue of both the femu and the lowe am. The thid objective integates esults fo the two main sets of analyses to detemine the effect of habitual behaviou, climate and body size on Neandethal long bone cuvatue. 3

35 1.2. Long bone cuvatue Fundamental to the study of skeletal chaacteistics, such as long bone cuvatue, is the hypothesis that the taits unde investigation ae functionally elevant and optimise mophology (Chuchill, 2005). The study of postcanial mophology ove the past decades has demonstated that skeletal mophology is unde vaiable envionmental and genetic influences. Theefoe, some featues give moe infomation about the biomechanical envionment (eason and Liebeman, 2004) while othes may yield moe infomation about the evolutionay histoy of a specific population (Ruff et al., 1991; eason, 2000a, 2000b; Liebeman et al., 2001; eason and Liebeman, 2004). The obseved vaiation in long bone cuvatue within and between species needs to be investigated using an appoach that consides its possible adaptive benefits. Long bone cuvatue is a complex featue to quantify, and its biomechanical envionment is difficult to model, as it is subject to diffeent stains duing diffeent stages of the gait cycle (Lanyon, 1980; Les et al., 1997; Main and Biewene, 2004). In humans, not all cuved bones ae active duing the gait cycle (e.g. adius and ulna) and may be subject to othe stains and stesses than when the same skeletal element is involved in locomotion in mammals that ae not bipedal. Because of this complexity, it has been difficult to assess the biomechanical ole and functional significance of diaphyseal cuvatue and the functional diffeences between bones and between species. Hominoids have a lowe degee of cuvatue than othe quadupedal mammals because thei elatively longe limb bones would endue vey high bending stess wee they as cuved as those of othe mammals (Biewene, 1983; Swatz, 1990; Betam and Biewene, 1992; Richmond and Whalen, 2001). The evolutionay significance of long bone cuvatue in hominins has, to date, not been investigated. Within humans, howeve, a ange of vaiation in femoal cuvatue has been epoted (Ried, 1924; Genna, 1930; Stewat, 1962; Walensky, 1965; Gilbet, 1975, 1976; Tudell, 1999; Buns et al., 2002) and, theefoe, it is vey likely that vaying degees of cuvatue in humans seve to educe individual habitual stain levels and to optimise function duing habitual behaviou in a specific envionment. It is unclea if the habitual stain levels in the lowe am and femu ae elated and that cuvatue is theefoe a systemic featue. 4

36 1.3. Neandethals and moden humans Ealy moden humans diffe fom ecent moden humans in both canial and postcanial featues but Neandethals diffe fom ecent moden humans much moe. Neandethals have a suite of chaacteistic canial taits such as a ounded canial vault; lage bowidges, lambdoidal flattening and an occipital bun; a low and long canium; a juxtamastoid pocess; supainiac fossa; a etomola gap; a chinless mandible; a lage nose; and mid-facial pognathism (Boule and Vallois, 1952; Tinkaus, 1983a; Hublin, 1989; Stinge, 1992; Hublin et al., 1998). In contast to the numeous diffeences in the canio-mandibula anatomy of Neandethals and moden humans, thee ae only a numbe of postcanial diffeences that have been identified as species defining. Most of these postcanial chaactes have been intepeted as the esult of the Neandethal hype-pola body shape and muscula hypetophy (atte, 1955; Vlcek, 1961b; Rak and Aensbug, 1987; Tompkins and Tinkaus, 1987; Holliday and Tinkaus, 1991; Ruff and Walke, 1993; Ruff et al., 1993; Walke and Leakey, 1993; Ruff et al., 1994; Tinkaus et al., 1994; Vandemeesch and Tinkaus, 1995; eason and Gine, 1997; Chuchill, 1998; Tinkaus et al., 1998a; Tinkaus et al., 1998b; Tinkaus and Ruff, 1999b; eason, 2000b; Holliday and Ruff, 2001; Shackelfod and Tinkaus, 2002; Maj et al., 2003; Weave, 2003; Thompson and Nelson, 2005; Shackelfod, 2007). Some of these postcanial anatomical specialisations include: a long pubic amus; an anteioly placed sacum; shot distal limb segments; a long glenoid fossa and a dosal sulcus on the scapula; lage ound apical tufts on the finges; a thick femoal and tibial shaft; and lage knees (atte, 1955; Vlcek, 1961a; Rak and Aensbug, 1987; Tompkins and Tinkaus, 1987; Holliday and Tinkaus, 1991; Ruff and Walke, 1993; Ruff et al., 1993; Walke and Leakey, 1993; Ruff, 1994b; Tinkaus et al., 1994; Vandemeesch and Tinkaus, 1995; eason and Gine, 1997; Chuchill, 1998; Tinkaus et al., 1998b; Tinkaus and Ruff, 1999a; eason, 2000b; Holliday and Ruff, 2001; Shackelfod and Tinkaus, 2002; Maj et al., 2003; Weave, 2003; Thompson and Nelson, 2005; Shackelfod, 2007). Othe chaacteistic Neandethal postcanial featues include a long distal phalanx in the thumb; flat capometacapal joint of the thumb; low femoal neck-shaft angle; absence of a femoal pilaste/linea aspea; and a cuved femu and adius (Aiello and Dean, 1990; Chuchill, 1998; Fleagle, 1999; Tinkaus, 2006). 5

37 Some of these featues may be pimitive etentions in Neandethals (Tinkaus, 1981, 1983a), wheeas othes may be autapomophic taits (Howell, 1957; Tinkaus, 2006). The taxonomic value of some of these postcanial featues, such as cuvatue of the femu and adius, has not been established, although it has been suggested that some postcanial featues, such as a geate level of obusticity, the absence of a pilaste and low neck-shaft angles, ae pimitive etentions (Tinkaus, 1983a; Ruff et al., 1993; eason, 2000b, 2000a). ostcanially, compaed to Neandethals, ealy moden humans ae chaacteised by high statue, high bachial and cual indices (Boule and Vallois, 1952; Tinkaus, 2007) and educed levels of obusticity which may eflect thei Afican ancesty (Mellas and Stinge, 1989; Aiello, 1993; Stinge, 2000; Stinge, 2002; Tinkaus, 2005). At the same time ealy moden Euopeans exhibit some chaacteistics which have been consideed to be distinctive Neandethal taits (Boule and Vallois, 1952; Tinkaus, 2007). These chaacteistics include aspects of the neuocanium, basicanial extenal mophology, mandibula amus and symphyseal fom, dental mophology and size and aspects of the clavicle, scapula, metacapals and appendicula popotions (Tinkaus, 2007). To some, the pesence of these Neandethal featues and the association of Neandethals with Uppe alaeolithic style tools (d'eico et al., 1998; d'eico, 2003; Ahen et al., 2004; Mellas, 2004; Mellas et al., 2007) suppots the idea that when moden humans migated out of Afica and into Euope thee was hybidisation between Neandethals and ealy moden humans. The extent to which this hybidisation took place and whethe o not it is still appaent in human mophology and genetics is a highly debated topic (Boule and Vallois, 1952; Smith et al., 1989; Faye et al., 1993, 1994; Wolpoff, 1996; Wolpoff and Caspai, 1997; Wolpoff et al., 2000, Deacon, 1992; Kings et al., 1997; Ovchinnikov et al., 2000; Hawks and Wolpoff, 2001; Caamelli et al., 2003; Caoll, 2003; Hagelbeg, 2003; Klein, 2003; Ovchinnikov and Goodwin, 2003; Geen et al., 2006; Noonan et al., 2006). The majoity of the liteatue on moden human oigins is focused on canial, mandibula and dental taits. ostcanial anatomy has eceived less attention, although thee ae some excellent desciptions of elevant postcanial mateial (Boule and Vallois, 1952; atte, 1955; Heim, 1983; Rak and Aensbug, 1987; Walke and Leakey, 1993; Vandemeesch and Tinkaus, 1995; Holliday, 1997; eason, 2000a, 2000b; Shackelfod and Tinkaus, 2002; Weave, 2003; Steudel-Numbes and Tilkens, 2004; Chuchill, 2005; Thompson and Nelson, 2005; Shackelfod, 2007; Aiello et al., 1999). What is evident is that Neandethals have a suite of 6

38 chaacteistics which, consideed independently, may occu in moden human populations, but which, as a suite, set apat the Neandethals as a goup that is distinct fom moden humans. The focus of most of the ealie wok has been on the paticulaities of Neandethal featues athe than a means of undestanding the evolutionay and adaptive pocesses that led to thei distinctiveness o what led to the divesity within moden humans and thei distinctiveness fom ealie hominins. Using a compaative method to distinguish Neandethal mophology fom that of ecent moden humans is useful but only when seen in the context of evolutionay biology and adaptive histoy. Thee ae thee main extenal influences that need to be consideed when intepeting the functional meaning of cuvatue, which is known to show a wide ange of intaspecific vaiation in moden humans. The fist is the effect of body size on cuvatue, because mammals show positive allomety with cuvatue. Ruff et a.l (1997) poposed that Neandethals ae on aveage 30% lage than ecent humans and that ealy moden humans ae about 10% lage than ecent moden humans (Ruff et al., 1997). If cuvatue is elated to body mass, it is pedicted that Neandethals will have highe degees of cuvatue than both ealy and ecent moden humans. Within moden humans, populations with the highest body mass ae pedicted to be moe cuved than those with lowe body mass. The second influence that needs to be investigated is the effect of habitual behaviou on cuvatue. Moden humans and Neandethals most likely did not diffe in thei subsistence stategies and wee pobably both hunting and scavenging (Liebeman, 1989; Ba-Yosef, 2004; eason et al., 2006). Although thee may have been diffeences in thei hunting pactices (Maean and Assefa, 1999; Maean and Assefa, 2005; Speth and Tchenov, 1998), thei esouce acquisition and oveall wokload involved high activity levels, and this is appaent in the similaities in thei post-cania (Liebeman, 1989; Tinkaus et al., 1989) If cuvatue is a esponse to activity levels in human populations, it is pedicted that Neandethals, having high activity levels, will display simila levels of degee of cuvatue to ealy moden humans and othe hunte-gathees. Within moden humans, it is pedicted that individuals and populations with lowe activity levels will exhibit lowe degees of cuvatue. Thidly, it is necessay to conside the effect of climate on cuvatue. Many of the distinctive Neandethal postcanial featues ae the consequence of a hypepola body fom (Hublin, 1989; Ruff, 1991; Weave, 2003; Weave and Steudel-Numbes, 2005). If the epoted high degee of cuvatue in Neandethals is one of those cold-adapted chaacteistics, ecent human populations 7

39 fom highe latitudes would be pedicted to possess highe levels of cuvatue than those fom lowe latitudes. Neandethals, being epoted as hype-pola (Weave, 2003), would be pedicted to have a highe degee of cuvatue than any moden human population. Climatic adaptations in humans ae known to become genetic adaptations ove time. In Neandethals and moden humans alike, it is expected that if thee wee a stong effect of climate on cuvatue that this would have been established in the population genetically athe than only though individual ontogeny. Though the pocess of genetic dift and isolation, ove time the distibution of the vaiation in cuvatue may have become a featue that has taxonomic value. By identifying the taxonomic value of cuvatue it may be possible to hypothesize about the elationship between ealy moden humans and Neandethals. If Neandethals ae distinct in thei long bone cuvatue fom ealy moden humans, and ealy moden humans esemble ecent moden humans moe than they do Neandethals, (Tinkaus and Shipman, 1993; Chuchill, 1998; Golovanova et al., 1999; Weave, 2003; Yamanaka et al., 2005 but see Shackelfod and Tinkaus, 2002). This would suppot the hypothesis that Neandethals wee excluded fom the evolutionay past of moden humans. 8

40 1.4. Layout of the thesis The second chapte povides an oveview of human and Neandethal vaiation in femu and lowe am anatomy and thei biomechanical popeties. The chapte continues with a discussion of the possible factos influencing cuvatue and concludes by outlining the specific hypotheses and associated pedictions in ode to addess the fist objective descibed above. Chapte 3 descibes the mateials, methods and statistical appoaches used in this eseach and ends with the ode of analysis. Chapte 4 contains the esults of the analyses of long bone cuvatue in ecent moden humans. The esults of the femu ae pesented fist, followed by the esults fo the lowe am. The chapte concludes with a discussion of the vaiation in long bone cuvatue in moden humans and summaises the pedictions fo the analyses on Neandethals and ealy moden humans. The esults fo fossil populations ae pesented in Chapte 5. Finally, Chapte 6 discusses the esults and conclusions fom this study. 9

41 CHATER 2. CURVATURE HISTORY OF RESEARCH ON LONG BONE 2.1. Femu Compaative anatomy of the femu Hominins like Homo habilis, Homo eectus, Homo heidelbegensis and Homo neandethalensis ae emakable in the similaity of thei femoal mophology (Kennedy, 1983b, 1983a, 1984). This mophology includes anteo-posteio flattening of the shaft eflected in the vitual absence of a pilaste, low neck-shaft angle, medial convexity of the shaft, a vey low minimal shaft beadth (waisting) and a medially expanded cotex at the mid-shaft level (compaed to anatomically moden humans whee the cotex is thickest on the lateal side of the shaft). This esults in a moe distal cossove of the biomechanical axis with the shaft axis (Kennedy, 1983a; Aiello and Dean, 1990). Both Tinkaus (1993) and Kennedy (1983a, b) have suggested that the medial convexity of the diaphysis and low neck-shaft angles ae a esult of highe activity levels (Kennedy, 1983b, 1983a; Tinkaus, 1993 but see Czanetzki, 2000). They suggest that this high activity level causes the femu to be moe medially convex poximally and to develop a lage tansvese diamete at mid-shaft (Kennedy, 1983b, 1983a; Tinkaus and Ruff, 1999b). Moe ecently, howeve, eseaches have agued that these featues in Neandethals might be a seconday consequence of a cold-induced body fom, elated to wide hips and moe obust extemities caused by the inteaction between genetically detemined body popotions and the magnitude of mechanical stess duing ontogeny o the diect consequence of vaiation in elative body size in individuals with cold-adapted bodies (Ruff, 1995; Weave, 2003). A cold-adapted body fom and wide pelvis may also explain the geate degee of femoal cuvatue obseved in Neandethals. The wide pelvis may esult in diffeent angles of hip joint eaction foce elative to the femu and affect the neck-shaft angle and tosion as the head of the 10

42 femu would be aticulating in a moe lateal position than in anatomically moden humans (Ruff, 1995). If the iliac blades ae oiented diffeently, this may lead to a moe anteio o posteio oientation of the acetabulae. Altenatively, the wide pelvis may simply cause an inceased distance between the acetabulae. Both these cases may lead to highe degee of cuvatue in ode to attain a hominin valgus angle Intaspecific vaiation in femoal cuvatue. In addition to the liteatue on Neandethal femoal cuvatue (see Chapte 1: Intoduction) seveal studies have investigated diffeences in femoal cuvatue among and between human populations in the light of biomechanical adaptation and foensic science. Foensic anthopologists studied femoal cuvatue as it was suggested to be a valuable tool to distinguish ace in human emains (Stewat, 1962; Walensky, 1962, 1965; Gilbet, 1975, 1976; Tudell, 1999). Stewat (1962) demonstated that thee was a diffeence in the expession of anteoposteio cuvatue of the femu between Caucasians, Afican-Ameicans and Native Ameicans (Dakota). Femoal cuvatue was measued as subtense by placing the distal condyles on a flat suface (Figue 2-1) and aising the poximal end so that the maximum concavity (deepest point on the anteio suface) on both distal and poximal ends ae at the same level (the levelling point). Then the distance was taken fom the table to the most anteio side of the femu. The analyses showed that shaft cuvatue was most ponounced in the Native Ameicans and least ponounced in Afican-Ameicans and that Caucasians occupied an intemediate position. 11

43 Figue 2-1 Subtense method employed by Walensky (1962). Native Ameicans also showed a geate amount of tosion compaed to Afican-Ameicans and Caucasians, with Afican-Ameicans showing the least amount. Individuals with highe degee of tosion also displayed a lowe apex of cuvatue. The positive coelation between cuvatue and tosion (Stewat, 1962) was not investigated futhe in subsequent studies on femoal cuvatue. Stewat concluded that although femoal cuvatue does not, as a ule, distinguish between aces, a femu with a maked degee of femoal cuvatue combined with a low degee of tosion distinguished a lage popotion of the Native Ameicans fom the Caucasians and Afican-Ameicans who have a lowe degee of cuvatue with a high tosion angle (Stewat, 1962). Walensky (1965) confimed Stewat s sepaation of Caucasians, Native Ameicans and Afican- Ameicans when he included the Inuit (Figue 2-2). He concluded that cuvatue inceased with age and population-elated functional activity and that diffeences in postual habits contibuted to these acial diffeences in femoal cuvatue (Walensky, 1962, 1965). 12

44 Figue 2-2 An Afican-Ameican, Inuit and a Native Ameican femu (fom Walensky, 1965) showing inceasing amounts of cuvatue and loweing apices of cuvatue. In 1976, Gilbet conducted an investigation into the possible causal factos of femoal cuvatue in Caucasians, Native Ameicans and Afican-Ameicans (Gilbet, 1976). He expanded Stewat (1962) and Walensky s (1965) sample with seven additional Native Ameican goups epesenting both pe- and post-colonial samples and looked at thei postual habits in elation to thei cuvatue. When only the Noth Ameican Native Ameicans wee taken into account, togethe with the Afican-Ameicans and Caucasians, Stewat s techniques distinguished Native Ameicans fom Afican-Ameicans o Caucasians. Howeve, when he included Native South 13

45 Ameican samples, the two goups combined showed only slightly moe ponounced femoal cuvatue than Afican-Ameicans. The South Ameican femoa wee less cuved than those of Caucasians and Noth Ameican Natives. Gilbet (1976) concluded that femoal cuvatue was not such a useful tool in ace assessment and set foth to look into possible causal factos of the tait (Gilbet, 1975). One of the hypotheses Gilbet (1976) tested was the effect of the equestian foaging lifestyle of the Noth Ameicans of the South Dakota aea (Aikaa: two sites dating between AD) on femoal cuvatue, but he noted that the non-equestian communities had the same degee of cuvatue as the equestian ones. The possibility that the euvian Natives wee less bowed because they wee fom an ealie sample was efuted because a moe ecent sample fell within the same ange of vaiation as the ancient sample (Gilbet, 1976). As mentioned in the pevious section, vaiation due to climate was efuted when Gilbet noted that two goups, living in the same egion, showed two diffeent anges of cuvatue and that the Inuit, expected to have the most cuved femoa, wee identical to those Natives living in the South. Gilbet (1976) agued that thee was little vaiation in postual habits between the goups and theefoe could not suppot Walensky s hypothesis that femoal cuvatue depended on postual habits. Instead, he agued that femoal cuvatue was genetically based but emained plastic and was influenced by goss body weight athe than by tempoal, climatic, postual o equestian influences. He suggested that obese individuals have a moe anteio cente of gavity which esulted in geate cuvatue. He did not follow up on the elation between tosion and femoal cuvatue (Gilbet, 1976). imate long bones ae less cuved than the long bones of othe mammals. Although in most anthopoids bones thee is an incease in cuvatue with body size (Swatz, 1990), expeimental wok has shown that the ontogenetic development of bone cuvatue in mammals depends on nomal muscle activity and weight-beaing (Lanyon, 1980) and is not influenced by individual vaiation in body weight. Whethe this is the case in humans needs to be detemined. Tudell (1999) evisited ace assessment though measuement of anteio femoal cuvatue and concluded that by inceasing the numbe of measuements taken on the bones, it is possible to disciminate Afican-Ameicans and Caucasians (Tudell, 1999). Maximal, bicondyla and oblique length wee measued as wee the midshaft and subtochanteic diametes. The cuve 14

46 was measued along thee points as the distance fom a flat suface when the femu is positioned in hoizontal position and balanced on two blocks with the distal condyles both touching a suface (Figue 2-3) (Tudell, 1999). Figue 2-3 Tudell s method of measuing cuvatue by placing the femu on two blocks (Tudell, 1999). A disciminant analysis with coss-validation on a seies of standad femoal measuements and the thee cuvatue distances of individuals of known sex and age categoy (below o above 30 yeas) povided an aveage accuacy of ace detemination of 87.12% fo both left and ight femu. This study was esticted to Afican-Ameicans and Caucasians but illustates the advantage of taking moe detailed measuements (Tudell, 1999) and the need to study wide anges of human populations. The lack of concodance among the eseach esults pesented above demonstates that thee is a need to investigate the vaiability of femoal cuvatue among a geogaphically and behavioually vaied ange of populations Biomechanics acting on femoal cuvatue To push the body upwads, i.e. when walking uphill, muscle foces extend the hip and the knee. Thee of the hamsting muscles (semi-tendinosus, semi-membanosus, long head of the biceps femois) extend the hip but do not ceate a significant bending moment in the bone and load it in uniaxial compession (Figue 2-4) (Fost, 1967). The fouth hamsting muscle (shot head of the 15

47 biceps femois) adds a posteio bending foce to the femu. The gluteus maximus and the two gastocnemii apply bending stess that bend the femu so it is convex posteioly (Fost, 1967; Cistofolini et al., 1995; Duda et al., 1996; Lengsfeld et al., 1996; Duda et al., 1997; Duda et al., 1998; Tinkaus et al., 1999b; Shackelfod and Tinkaus, 2002; Hall, 2004). Figue 2-4 Muscles acting on the femu. l: pelvis. F : femu. :patella. T : tibia. SM : thee muscles; semitendinosus, semimembanosus, long head of the biceps femois. GM: gluteus maximus. GN: gastonemius. QF: quadiceps femois. A: The femu of a man walking up a step. Thee is a bending foce acting on the femu making it posteioly convex. B: SM ae thee of the fou hamsting muscles. They extend the hip and do not ceate bending moments but compession. The shot head of the biceps femois (BF) adds posteio bending. C: The gluteus maximus bends the femu so that is posteioly convex. The gastocnemii add to this bending foce. D: The quadiceps bends the femu in the opposite way. This dynamic inteacting muscle system minimises bending foces in the femu (afte Fost, 1967). 16

48 The quadiceps muscles exet stess on the femoal shaft in the opposite diection than the gastocnemius, shot head of the biceps femois and the gluteus muscle so that the shaft is anteioly convex, ceating a balance in the muscle foces acting on the diaphysis. This balance minimises the bending stesses on the femu (Fost, 1967). In most quadupeds, this balance is close to pefect and femoa show little o no anteoposteio cuvatue in the diaphysis. In humans thee is a esidual anteo-posteio bending visible (Fost, 1967). 17

49 2.2. Radius and ulna Compaative anatomy of lowe am anatomy Fom the well ponounced muscle aticulations on all uppe limb bones, it is suggested that Neandethals had vey poweful foeams (Tinkaus and Chuchill, 1988). Thee ae featues in the ulna and adius that distinguish Neandethals fom moden humans (Fische, 1906; atte, 1955; Tinkaus and Chuchill, 1988; Aiello and Dean, 1990; Vandemeesch and Tinkaus, 1995; eason and Gine, 1997). The poximal ulna is diffeent in that the tochlea notch is oiented moe anteioly in Neandethals than it is in moden humans. Tinkaus and Chuchill (1988) popose that this would not have limited the ange of movement but was athe an expession of diffeent habitual behaviou such as the inceased use of foeams with the elbow flexed. The ponato quadatus cest is vey ponounced and also suggests a moe muscula foeam, although the inteosseous cest is pooly developed and the shaft is elatively naow (Tinkaus and Chuchill, 1988; Aiello and Dean, 1990). The supinato cest is stongly developed and the shaft shows a geate degee of lateal cuvatue than that found in moden humans. This may indicate that Neandethals closely esemble ealie hominins in the mophology and stength of the adius and that the Neandethal foeam and elbow was especially stong duing ponation and supination (Tinkaus and Chuchill, 1988). The position of the adial tubeosity is a measue of leve advantage of the biceps bachii. In the apes, it is positioned moe medially. This gives apes a geate mechanical advantage of the biceps bachii in supination. The tendons wap themselves aound the adial shaft and the medial position of the insetion and inceases the distance between the poximal and distal insetion of the muscle and esults in a lage medial otation axis of the foeam. If the adial tubeosity is placed moe anteo-lateally, as it is in moden humans, then powe advantage is lost duing the 18

50 final phases of supination (Tinkaus and Chuchill, 1988; Aiello and Dean, 1990; eason and Gine, 1997). The adius cuves mainly in a medio-lateal plane while the ulna tends to cuve in a dosovental plane. A geate distance between them inceases the distance between the insetions of the ponato quadatus and the ponato tees. Afican apes ae less cuved than othe mammals. Swatz (1990) suggests this is due to long bones of pimates being longe than those of othe mammals and will theefoe poduce lage bending stesses duing nomal locomotion. Highe degees of adial cuvatue in anthopoids have been explained to be the esult of an incease in size and functional impotance of the supinato musculatue, but in gibbons was not affected by diffeential muscle mass (Swatz, 1990). Compaed to humans howeve, apes have a highe degee of lateal cuvatue. The highe degee of cuvatue in Afican apes (Matin and Salle, 1959; Knussman 1967 in Swatz, 1990) and a moe lateal insetion of the ponato tees inceases the leve advantage (Aiello and Dean, 1990). Figue 2-5 Hominoid adii. Right adii of A=Goilla, B=an, C=ongo, D= La Chapelle-aux-Saints, E= La Feassie I, F= La Feassie II, G= ecent Euopean (Afte Czanetzki, 2000). 19

51 The lateal subtense of the adius of the Neandethals is emakable and falls on o beyond the highe limits of the moden human vaiation (Fische, 1906; Botez, 1926 in atte, 1955; Vandemeesch and Tinkaus, 1995; Caeteo et al., 1999; Czanetzki, 2000) (Figue 2-5). Although some confusion exists about which technique yields the most accuate measuement of cuvatue of the lateal side of the adius (See Matin and Salle, 1959 fo fou diffeent methods to measue cuvatue), only Fische (1906) epots a size coected measue o an index of cuvatue (subtense/maximum length*100). Quantification of the posteio cuvatue of the ulna using a subtense technique (Fische, 1906; Matin, 4a) is not as staightfowad as it is fo the moe evenly shaped bones such as the femu and the adius, but Neandethals have been descibed as having a lage posteio subtense in the ulna (Fische, 1906). The head/length atio of the adius (head diamete/length*100) is lage in the Neandethals than it is fo any othe human population, but thee is a lage ange of vaiation within moden humans (atte, 1955). Fische (1906) and atte (1955) also epot an enlaged distal condyle fo the Neandethals and comment on the pesence of this enlagement in Japanese, Aficans, Austalians and othe human populations. The enlagement of the condyles may be caused by the otation of the adius (Fische, 1906), but atte (1955) wans that this may not be as staightfowad in hominins as in mammals whee thee is a elation between otation and size of the condyles. He also wans biomechanics is not always the cause of lage condyles but that they have also been associated with ickets (Mafan, 1912 and Decugis, 1941 in atte, 1955; Steinbock, 1976; Ivanhoe and Tinkaus, 1983). Investigations of the ulna and adius have shown that ealy anatomically moden humans have elatively thick cotical bone compaed to ecent moden humans (Chuchill et al., 1996; eason and Gine, 1997; Gine et al., 1998; eason et al., 1998) and that ealy moden humans have a ticke and shote adial neck than Neandethals (Chuchill et al., 1996; eason and Gine, 1997) Intaspecific vaiation in the adius and ulna Thee ae vey few studies on vaiation in longitudinal cuvatue of the adius and ulna within moden humans. A summay of the mophological vaiation in moden humans in the ulna and adius is descibed below. 20

52 Within ecent human populations, the distal limb segments (tibia and adius) tend to exhibit moe elative vaiability (size independent) than the poximal segments, especially in the lowe limb (Holliday and Ruff, 2001). Males and females appea to be slightly diffeent in this patten. Females vay to an equal degee in both uppe and lowe distal segments, wheeas males show most vaiability in the lowe limb (Holliday and Ruff, 2001). These diffeences ae believed to be allometic since males ae lage than females and this allomety can also be found when looking at between-goup diffeences in, fo example, cold- and wam-adapted populations (Holliday and Ruff, 2001). Reseach on ecent human vaiation of the ulna and adius is limited and most of it dates back to the ealy 20 th Centuy. In 1906, Fische made an in-depth study of the vaiation of the adius and ulna and included both Neandethal casts and ecent moden human populations fom diffeent geogaphic oigins. His sample of moden humans consisted of Euopeans, Aficans, Austalians, olynesians, Melanesians, Bimese, Tieo del Fuegans, Ainu, Japanese, hilippinos and pehistoic Egyptians. atte (1955) included this study and othes in his book on Neandethals and summaised some of the main diffeences between moden humans and Neandethals. Lapps, Japanese and Medieval Euopeans have moe obust adii than do Neandethals. The Aficans have the smallest obusticity index but thee is a lage amount of vaiation. Also, the Neandethal ulna is obust fo its size (Fische, 1906; atte, 1955). Fische (1906) epots a mean index fo humans in lateal subtense of the adius anging fom 2.5 fo the Tiea del Fuegans to 3.2 fo the Euopeans compaed to a mean of 7.4 (S.D.=2.5, n=5, summay data fom Caeteo et al., 1999) fo the Neandethals. Klaatsch (1901) suggests that adial cuvatue is a heeditay tait. Howeve, because humans ae geneally bon with staight ulnae and adii, Rouvièe (1939, in atte, 1955) agues that adial lateal cuvatue is a biomechanical adaptation to the stong development of the flexo muscles of the finges and thumb. The mediolateal cuvatue of the anteio suface of the ulna is difficult to descibe because of the sinusoidal shape of the diaphysis. Fische (1906) used diaphyseal angles fo each cuve in the anteio ulna and found that Euopeans ae the least cuved, and that Austalians and Tiea del 21

53 Fuegans possess the highest degee of cuvatue. atte (1955) does not epeat this method and does not comment on the cuvatue of the ulna in Neandethals. Fische (1906) measued the angle the adial tubeosity makes with the pependicula plane though the styloid pocess and the ulna notch. This measuement will give the angle that the adial tubeosity deviates fom the axis though the inteosseous cest. Moden humans ange fom 0 to 85 (Fische, 1906) with the majoity anging between 45 and 60 (Boule and Vallois, 1952). Thee is a lage ange of vaiation within single populations with angles. Fo example, Euopeans ange fom (mean=50.2 ) and Aficans fom (mean=63.3). Although a vey high angle (fom 81 Spy 1 88 Neandethal) was consideed a deived Neandethal featue, in moe ecent papes, the angle of the adial tubeosity is appoximated qualitatively, and it was concluded that although Neandethals have a vey high angle and theefoe a moe medially oiented adial tubeosity, they do not fall outside of the ange of vaiation of moden humans (Tinkaus and Chuchill, 1988; Vandemeesch and Tinkaus, 1995). Fische (1906) suggests a coelation between the length of the biceps bacchii muscle tendon and the position of the adial tubeosity. When the am is pat flexed in ponation, with the hand in supination o semisupination, thee is a stain on the biceps and theefoe the tendon and the tubeosity moves. Habitual use of the am in that position can cause the individual diffeences obseved in the oientation of the adial tubeosity (Fische, 1906; Tinkaus and Chuchill, 1988; Aiello and Dean, 1990). When adial neck length is coected fo size by the adial length, the Neandethals have a elatively long adial neck fo adial length and fall with the Aficans and Chinese athe than with the Euopeans (Vandemeesch and Tinkaus, 1995). A longe adial neck makes the biceps bachii moe effective as it has moe leve advantage and theefoe geate powe. Thee is a lage ange of vaiation in adial neck-shaft angle within moden human populations but the Euopeans have been suggested to have the lagest when compaed to othe populations (Fische, 1906). The joint-axis angle (o neck-shaft) of the ulna is the angle the tochlea notch makes with the shaft axis and is measued by finding the angle between the sagittal axis of the tochlea notch and the shaft axis. In humans, it vaies between 0 and 28 and Austalians, hillipinos and 22

54 Tiea del Fuegans have slightly highe angles, but thee is no tend among populations and no coelation between angle and cuvatue was obseved (Fische, 1906). Vey few studies have exploed the behavioual and envionmental factos on lowe am mophology. Robusticity of the uppe limb, howeve, has been investigated in elation to climate and habitual behaviou (Stock, 2002; Stock and feiffe, 2004; Stock, 2006). Although climate has a significant influence on pattens of diaphyseal obusticity, pattens of obusticity of the uppe limb coespond best to maine mobility especially in the distal limb elements. This suggests that thee may be geate diaphyseal plasticity futhe away fom the tunk and that diffeences in bone mass in the lowe am ae moe elevant fo functional intepetation of achaeological and fossil samples without being constained by the enegetics of bipedal locomotion (Stock, 2002; Stock and feiffe, 2004; Stock, 2006) Biomechanics acting on lowe am cuvatue The elbow joint acts as a leve and is composed of the humeo-ulna, humeo-adial and poximal adio-ulna joints. All ae encapsulated by collateal ligaments. The humeo-ulna joint is composed of the tochlea that aticulates with the tochlea fossa of the ulna. This joint seves in flexion and extension. The humeo-adial joint is lateal to the humeo-ulna joint. It is fomed between the distal pat of the humeus and the head of the adius. This joint is not fixed but is esticted in its movement by the humeo-ulna joint. It is used duing flexion, extension, supination and ponation. In the poximal adio-ulna joint, the head of the adius aticulates with the adial notch of the ulna. This joint pivots duing ponation and supination making the adius oll ove the ulna in a medial and then lateal fashion (Fost, 1967; Hall, 2004). The lage numbe of muscles poducing the ange of motion of the elbow and foeams complicates a foce-analysis fo this complex of joints. It is assumed, howeve, that the stongest flexo muscle of the elbow is the bachialis. Distally, bachialis insets below the coonoid pocess. Anothe elbow flexo, the biceps bachii, insets in the adial tubeosity and is stongest duing supination. The bachio-adialis also aids in flexion and is most effective in the neutal position (between ponation and supination). Its distal insetion is in the base of the styloid pocess on the lateal aspect of the adius. The stongest extenso muscle is the ticeps. The thee heads of the ticeps inset on the olecanon pocess of the ulna with a common tendon. The 23

55 anconeus muscle attaches to the lateoposteio aspect of the ulna and is only a mino extenso of the elbow (Fost, 1967; Hall, 2004). The ponato tees muscle, the supinato and the ponato quadatus ae involved in ponation and supination. These ae inseted in the poximal and distal adio-ulna joints. The inteosseous space between ulna and adius detemines the degee of ponation and supination an individual can achieve (Yasutomi et al., 2002). Yasutomi (2002) used thee dimensional models to econstuct diffeent sizes of inteosseous space and found that when the axis of otation in ponation and supination passed though the inteosseous egion the otation was moe than 40% adially, ulnaly, anteioly and posteioly. Howeve, when the axis of otation was deviated fom this egion, thee was significant loss of supination and ponation (14% adially, 7% ulnaly, 5% anteioly and 4% posteioly) and estiction by the elastic inteconnecting membane (Yasutomi et al., 2002). The ponato quadatus is the majo ponato muscle and is assisted by the ponato tees. The ponato quadutus attachments ae on the lowe anteio ulna and the lowe anteio adius. The ponato tees insets lateally in the middle of the shaft of the adius and has a mino ole in flexion. The supinato muscle is the majo supinato and is assisted by the biceps when the elbow is flexed to 90 o less. The supinato muscle insets on the lateal poximal pat of the ulna and the lateal poximal pat of the adius (Hall, 2004). The elbow is not a weight-beaing bone but sustains lage loads thoughout its activity cycle. Most of the compessive loading is at the elbow and geate foces ae geneated when the hands ae otated in ponation. Lage foces ae also geneated duing cetain activities. As the attachment of the ticeps muscle on the ulna is close to the elbow than ae the bachialis and the biceps, the moment am is smalle and because of this leve advantage, the flexo muscles have to geneate less foce than the extensos to ceate the same amount of joint toque (Fost, 1967; Hall, 2004). 24

56 2.3. ossible causes fo vaiation in long bone cuvatue Neandethals and ickets Some scholas have suggested that Neandethal cuvatue in the ulna and adius is the esult of ickets (Ivanhoe, 1970; Ivanhoe and Tinkaus, 1983; Czanetzki, 2000) o osteomalacia (Czanetzki, 2000). Rickets is a medical condition wheeby the osteoid (the oganic mateial in bone) fails to calcify in a gowing animal o human. Individuals with ickets have a deficient vitamin D metabolism. Othe dietay deficiencies in the calcium o phosphous metabolism may poduce ickets. This esults in skeletal defomity and shot statue. Figue 2-6 X-ay image of an infant with sevee ickets. Note the medio-lateal cuve as opposed to the the anteo-posteio cuve obseved in Neandethals. Fom (last accessed 19/06/2008) 25

57 Natual vitamin D is fomed in the skin unde the stimulus of ultaviolet light and is pesent in fish live oil (Stuat-Macadam and Iscan, 1989; Wood et al., 1992). Because thee is no widespead evidence of Neandethals eating fish (with the exception of shell fish consumption at Gibalta) (Hockett and Haws, 2005) and thei inhabitation of the Nothen egions of Euope, Ivanhoe suggests Neandethals expeienced an insufficient amount of vitamin D in thei diet and as a consequence of ickets show skeletal defomities such as abnomal long bone cuvatue (Ivanhoe, 1970; Ivanhoe and Tinkaus, 1983; Czanetzki, 2000). Howeve, the cuvatue obseved in Neandethals is an accentuation of nomal anteoposteio cuvatue of the diaphysis (Steinbock, 1976) and neve assumes the iegula mediolateal cuvatue associated with ickets (Figue 2-6) (Ivanhoe and Tinkaus, 1983). Neithe does ickets explain the obseved vaiation in anteio cuvatue between moden human populations Biomechanics and bone emodelling Wolff s Law states that bones gow and emodel thoughout an individual s life in ode to adapt to thei mechanical envionment. The bone senses, tansduces, and esponds to loads by molecula and physiological mechanisms (eason and Liebeman, 2004; Ruff et al., 2006). Long bones of othe teestial mammals also display some longitudinal long-bone cuvatue and the magnitude of this may vay acoss bones, species and even between individuals (Lanyon and Baggott, 1976; Lanyon, 1980; Biewene, 1983; Lanyon, 1987; Swatz, 1990). Seveal mammals have been used in expeimental studies to investigate the functional meaning and development of longitudinal cuvatue and how this may affect stain and stess distibutions in the shaft (Fost, 1967; Lanyon and Boun, 1979; Lanyon, 1980; Biewene, 1983; Lanyon, 1987; Betam and Biewene, 1988; ead and Lanyon, 1990; Swatz, 1990; Les et al., 1997; Main and Biewene, 2004; Yamanaka et al., 2005). Seveal studies (Lanyon, 1980; Biewene, 1983; Betam and Biewene, 1988; Betam and Biewene, 1992; Biewene and Betam, 1994; Main and Biewene, 2004) have established that if thee is an absence of loading fom muscle activity and weight-beaing duing ontogeny, long bones fail to develop thei appopiate bone mass o longitudinal cuvatue, despite achieving thei nomal length. Lanyon (1980) concluded that thee ae cetain aspects of bones that ae genetically detemined but that othe featues equie a nomal mechanical envionment to develop. 26

58 Liebeman and eason (2001) pefomed an expeimental study testing the hypothesis whethe cotical bone gowth (modelling) and epai (Havesian emodelling) ae esponses to execiseinduced mechanical loading and whethe the emodelling vaied with loading and the position in the skeleton. Execised juvenile sheep had highe peiosteal modelling than Havesion emodelling ates than non-execised contols (Liebeman and eason, 2001). Mid-shaft peiosteal gowth was highe poximally and mid-shaft Havesian emodelling was highe distally. Gowing animals thus modulate modelling vesus emodelling,espectively,to loading at diffeent skeletal locations. This is to optimize coss-sectional stength elative to the kinetic enegy cost of acceleating added mass (Liebeman and eason, 2001). Ruff et al. (2006) suggest that ates of emodelling and ates of bone tunove vay geatly at diffeent skeletal sites and that thee is no simple elation between the oientation of loads, such as stains and stesses, and the coss-sectional geomety of long bones (Liebeman and eason, 2001; Ruff et al., 2006). If, howeve, cuvatue only develops unde a nomal developmental activity egime, it can be assumed that it has a functional advantage to eithe the bone itself o to the anatomical stuctues aound it. The elationship between foces and modelling and emodelling of long bones is complex. If one consides the long bone as a long and slende beam, it is assumed that the optimal function of this bone to esist applied stesses and minimise stain is though axial compession (Fost, 1967; Betam and Biewene, 1988; Hall, 2004). This loading configuation distibutes most mateial in the plane of defomation, and cotical bone is stonge unde compession than unde tension (Fost, 1967; Lanyon and Baggott, 1976; Lanyon, 1980; Betam and Biewene, 1988; ead and Lanyon, 1990; Hall, 2004). Applying axial loading to a bone that is longitudinally cuved, esults in a bending moment that is popotional to the displacement of the diaphysis pependicula to the longitudinal inteaticula axis (Fost, 1967; Swatz, 1990; Hall, 2004). Because of this bending, tensile and compessive stesses ae unevenly distibuted though the bone and even small extenal loads can ceate lage stains within the bone (Lanyon, 1980). Reducing cuvatue while axially loading long bones should esult in the lowest stain levels. Howeve, the long bones of mammals ae not loaded puely axially and long bones can expeience significant bending moments due to cuvatue and muscle and joint eaction foces that ae not pefectly aligned with the axis of the bone (Betam and Biewene, 1988). Also, in 27

59 the human femu, the positions of the aticulation and muscle attachments, such as the medial displacement of the femoal head (Tinkaus, 1993; Andeson and Tinkaus, 1998), and the contaction of the adducte and gluteal abducto muscles, cause the femu to be subjected to some degee of mediolateal bending (Ruff, 1995). In one legged stance, most of that bending stess may be educed though associated tension in the iliotibial tact and musculatue (Lengsfeld et al., 1996; Taylo et al., 1996; Les et al., 1997; Simões et al., 2000). Ruff (2000) suggests that anteoposteio bending could be the cause of anteoposteio expansion of the femoal midshaft in esponse to high activity and mobility levels (Ruff, 2000b). Taylo and colleagues (1996) investigated loading though the femu in one-legged stance in humans by measuing the dominant mode of loading in the femu in a finite element analysis. In a finite element analysis the mateial popeties and loading of the skeletal elements o joints ae modelled and analysed to bette undestand the biomechanics and othopedics (Richmond et al., 2005). The esults showed that the human femu is loaded pimaily though compession athe than though tosion o bending (Taylo et al., 1996). They also found that the anteio and posteio stesses on the femu ae negligible and that this is pobably due to the eduction of oveall bending stesses in the femu due to the application of muscle foces. If a bone is loaded in bending, this would incease the biological and locomoto cost of bone poduction because the bone would need to esist these stesses and consequently be thicke (Taylo et al., 1996; Skey, 2008). When the femu is loaded though bending stess, one would expect deflection of the femoal head and an uneven load tansfe though the distal condyles but evidence shows unifom pessue distibution in both condyles (Taylo et al., 1996). The majo limitation of this study was that it was done duing one phase of gait and theefoe is not necessaily applicable to the whole gait cycle. It may be that the loading stesses diffe thoughout the cycle but anatomical featues of the femu suggest this is not so. Duda et al. (1996) found that diffeences in muscle attachments esult in diffeent biomechanical popeties of individuals. Not only is bone emodelled when applying diffeent stesses, but so ae the soft tissues such as muscles and tendons. Duda et al. also ecognise that when one neglects the majo muscles, compession, bending and tosion may be oveestimated and not play as significant a ole as fist assumed and etun the diaphyseal bending stesses to ones of axial compession (Duda et al., 1997; Duda et al., 1998). 28

60 Modelling the system of inteacting muscles and bone in stance is impotant fo undestanding the functional significance of cuvatue but does not explain diffeences in femoal cuvatue between individuals as it emains difficult to measue the in vivo levels and distibutions of diaphyseal stains in individuals (edesen et al., 1997). Also the complex ways in which muscles o pats of muscles contact and of joint eaction foces duing gait with vaying budencaying levels in a natual setting, make it impossible to fully undestand the esulting stains in the femu, especially as thee is evidence fo vaiation in the human femoal muscles that would cetainly affect the muscle foces applied to the femu (Duda et al., 1996). Nonetheless, it emains possible that cuvatue seves to lowe bending stesses elative to staight bones by educing bending moments placed on the diaphysis and in that way etuning the bone to an envionment of axial compession (Fost, 1967; Hall, 2004). Most expeimental wok though, has demonstated that cuvatue inceases bending stains and that the diection of the cuve does not necessaily coespond with the tension suface of a bone when it is loaded (Lanyon and Baggott, 1976; Lanyon and Boun, 1979; Lanyon et al., 1979; Lanyon, 1980; Biewene, 1983; Lanyon and Rubin, 1986; Lanyon, 1987; Swatz, 1990; Simões et al., 2000). Fo a weight-beaing bone, longitudinal cuvatue may be cucial because it educes the ability to withstand high levels of loading and be a compomise between bone stength and pedicting bending stains and mateial failue (Lanyon, 1980, 1987; Betam and Biewene, 1988). Betam and Biewene (1988) ague that axial compessive loading is unstable as a catastophic shift fom compessive stess to bending stess in a staight column is equally likely to bend in a andom diection. A cuved bone, howeve, is moe likely to bend in the diection of its longitudinal cuvatue egadless of the oientation of the bending moment applied to the bone and is theefoe pedictable. Alexande (1981) demonstated that stuctues that ae likely to be subjected to unpedictable loads would need to build in a safety facto fo maintaining the biological stuctue, even if that safety facto would be moe metabolically costly to maintain and tanspot (Alexande, 1981). The final anatomy of the bone will thus be a compomise between the demands of load caying (cuvatue negatively affects stength) and pedictability (Betam and Biewene, 1988). Lanyon and Boun (1979) also suggest that femoal bending may facilitate lage muscle packing and/o place the muscle vecto moe paallel to the diaphyseal axis. Cuvatue allows fo the positioning of lage muscle bellies while allowing the slende muscle attachments to be close 29

61 to the joints. Having muscles adjacent to the bone exets pessue on the peiosteum, inceases bone esoption, and may cause cuvatue. This hypothesis is suppoted by the concavity of the adius and tibia of many mammals with espect to the flexo musculatue, allowing fo geate volume (Lanyon et al., 1979; Lanyon, 1980). The pesence of intemediate stains fom cuvatue-induced bending stess may also be advantageous fo bone to maintain a minimum bone mass. Reduction of loading esults in a decease in bone mass (Lanyon and Baggott, 1976; Lanyon and Boun, 1979; Lanyon et al., 1979; Lanyon, 1980; Ruff et al., 1991; van De Meulen et al., 1993; Cate et al., 1996; Liebeman et al., 2001; Liebeman and eason, 2001; eason and Liebeman, 2004; Ruff et al., 2006). Theefoe, if the bone was loaded in puely axial compession, thee may not be enough stain fo the bone to benefit physiologically. Stain levels can be inceased by augmenting the degee of bone cuvatue o by educing bone coss-sectional aea and/o second moment of aea until an optimum between physiological benefit and isk of failue has been achieved (Lanyon, 1980). To summaise, thee ae fou main biomechanical hypotheses explaining longitudinal cuvatue of the long bones: 1) cuvatue lowes bending stess by tanslating bending stess to axial compession (Fost, 1967; Hall, 2004), 2) cuvatue facilitates muscle expansion and packing (Lanyon et al., 1979; Lanyon, 1980), 3) cuvatue is a compomise between bone stength and pedictability of bending stains and mateial failue (Lanyon, 1980, 1987; Betam and Biewene, 1988), o 4) geneates stains necessay fo optimal stength (Lanyon, 1980). 30

62 Body size It is undestood that loading of the long bone diaphysis is popotional to body mass (Ruff, 2000b). Robusticity, which is a esponse to loading, has an allometic elationship with body size (van De Meulen et al., 1993; Ruff, 2000a; Stock, 2002; Stock and feiffe, 2004). Anthopoids show and oveall positive allomety in thei cuvatue (Swatz, 1990) so lage anthopoids have a highe degee of cuvatue. This allometic elationship is simila to that of a boade goup of mammals (Swatz, 1990 but see Biewene, 1983; Betam and Biewene, 1992) but pimates ae much less cuved than mammals at any given size in ode to allow fo elatively longe limbs but etaining low levels of bending stess (Swatz, 1990) Activity levels Vaiation in obusticity levels is often suggested to be an adaptation to activity levels and habitual behaviou, and a substantial amount of eseach has focused on the changes in skeletal obusticity thoughout human evolution and the evidence fo oveall gacilisation (Ruff et al., 1993; Ruff et al., 1994; Tinkaus et al., 1994; Tinkaus et al., 1999a; eason, 2000a, 2000b; Ruff and Tinkaus, 2000; Shackelfod and Tinkaus, 2002; Shackelfod, 2007). Seveal ecent studies have also been conducted to undestand pattens of skeletal obusticity in moden humans (Lasen, 1995; Ruff and Tinkaus, 2000; Stock and feiffe, 2004; Stock, 2006; Calson et al., 2007). Undestanding pattens in obusticity may aid in undestanding long bone cuvatue if both ae emodelling esponses to simila stains and stesses. The elationship between skeletal obusticity and habitual behaviou, and moe specifically teestial mobility, has been investigated pimaily using mid-shaft femoal coss-sectional geomety. This eseach is based on the pediction that epetitive anteoposteio loading on the lowe limb duing subsistence stategy-elated teestial mobility will esult in thickening of the coss-sectional geomety in the anteoposteio plane (Ruff, 1987, 1994a; Lasen et al., 1995; Holt, 2003; Stock and feiffe, 2004), and this is suppoted by the stength ciculaity indices (I x/ I y ) at the femoal midshaft and its stong coespondence with teestial obusticity (Stock, 2006). If thee is a coelation between obusticity and cuvatue, the anteoposteio bending that is obseved may be a esponse to the inceased cuvatue of the diaphyseal shaft. 31

63 Holt (2003) demonstates thee is a elationship between femoal anteoposteio bending stength, lowe limb obusticity and declining teestial mobility fom the Uppe alaeolithic though Mesolithic in Euope. Changes in postcanial obusticity with a shift away fom hunting and gatheing and the adoption of agicultue also suggest that inceased sedentism is visible in the extenal (Ruff et al., 1984; Lasen, 1995 but see Bidges, 1989a; Bidges et al., 2000) and intenal dimensions of long bones (Ruff, 1987; Bock and Ruff, 1988). This is suppoted by the highe pevalence and seveity of osteoathitis in hunte-gathees compaed to agicultualists (Otne, 1968; Lasen, 1983; Bidges, 1989b; Lasen, 1995). Although this patten of deceasing obusticity is pesent in human populations, geneally, males appea to be moe ponounced in thei eduction than females. This may eflect the changes in types of activity that wee geate in males than they wee in females (Ruff, 1987). This compaison of coss-sectional geomety and the anteio-posteio bending stess (I x ) and medial-lateal bending stess (I y ) is accompanied by a eduction in sexual dimophism with the tansition fom hunting to gatheing to agicultue (Ruff, 1994a). Sexual dimophism in hunte-gathees is the esult of the ole of males to tavel long distances and hunting compaed to the moe sedentay ole of females in gatheing and childcae (Ruff, 1987). Recently, obusticity has been investigated thoughout the skeleton and thee is a gowing body of evidence that aquatic foaging and the habitual use of watecaft fo subsistence has an influence on uppe limb obusticity (Stock and feiffe, 2001; Weiss, 2003; Stock and feiffe, 2004; Stock, 2006; Shackelfod, 2007). Thee is a tend fo distal elements to show a stonge elationship between hypetophy and behaviou but obusticity at femoal midshaft (measued as stength ciculaity index shape index) shows the geatest coespondence to teestial mobility. Recently, it has become inceasingly clea that the elationships between postcanial obusticity, mobility and activity pattens ae not as staightfowad as initially believed and that levels of obusticity may vay at diffeent sites of the bone (Stock, 2006). In the limbs, coelation between obusticity and teestial o maine mobility inceases fom poximal to distal. Theefoe, stonge elationships would be expected between bone modelling and emodelling in esponse to stain in the distal elements compaed to poximal elements (Stock and feiffe, 2001; Stock, 2006). 32

64 Diaphyseal obusticity in the uppe limb bones have often been used as evidence fo diffeences in habitual behaviou thoughout human evolution (Tinkaus et al., 1994; Vandemeesch and Tinkaus, 1995; eason et al., 1998; Tinkaus et al., 1999a; eason, 2000a, 2000b; Ruff and Tinkaus, 2000) and Stock (2006) suggests that thee is geate vaiability in the obusticity of the distal limb segments that is associated with habitual behaviou, especially in the mid-shaft of the ulna. If long bone cuvatue is a esponse to activity levels and habitual loading, it is pedicted to be highest in populations with high activity levels (Ruff et al., 1984; Lasen, 1995; Ruff, 1999) and to vay between males and females (paticulaly in hunte-gathees) (Bock and Ruff, 1988; Ruff, 1994a; Lasen, 1995). Also, with inceasing sedentism though time, a deceasing degee of cuvatue would be pedicted. The complexity of the elationship between loading and obusticity is subject to additional factos, the main ones being the susceptibility of bone to stain duing ontogeny (Ruff et al., 1994; Liebeman et al., 2001; eason and Liebeman, 2004) and the effect of climate (eason, 2000b; Weave, 2003) Climate Climate affects body size and popotions and it has been suggested that geate obusticity in individuals fom colde climates may be an indiect effect of a lage body size (Tinkaus and Ruff, 1999b; Tinkaus and Ruff, 1999a; Stock, 2006). Othe studies have found a diect effect of climate on coss-sectional geomety (Stock, 2006) and extenal obusticity (Ruff, 1995; eason, 2000b; Weave, 2003; Stock, 2006). Begmann and Allen s ules apply to body size and popotions in mammals and thei elation to themo-egulation. Thee is a positive elationship between body size (weight) (Begmann, 1847) and a negative elationship between limb length elative to body mass with inceasing distance fom the equato (Allen, 1877). Consideable studies on a ange of human populations have confimed these pinciples also apply to humans. Body beadth is coelated most stongly with tempeatue, and diffeences in limb popotions and body size ae established though 33

65 genetic adaptation and not though individual ontogeny (Y'Edynak, 1976; Eveleth and Tanne, 1990; Ruff et al., 1994; eason, 2000b; Van Andel, 2003; Weave, 2003; Ruff et al., 2005). Thee have been ecent changes in the compliance of moden humans to these ecological pinciples due to dietay impovements of many hunte-gathees and the adoption of a moe uban tading subsistence stategy (Katzmazyk and Leonad, 1998). Theefoe, cae must be taken when analysing diffeences within moden humans and especially when dawing conclusions fo palaeoanthopological studies (Stock, 2002). In an evolutionay context, body size and limb popotions have been used to intepet envionmental adaptation and migation, especially when explanations ae sought fo the diffeences in Neandethal and ealy moden human body build. Weave (2003) agued that the elationship between obust femoa and cold climate in Neandethals can be explained as a seconday consequence of the wide cold-adapted Neandethal bodies and that the shape of the Neandethal femu can be explained as a seconday consequence of the cold-adapted bodies vs. the wam adapted bodies of moden humans (Weave, 2003).. Because the beadth of the pelvis is much wide in Neandethals, the femu esponds to this with lage aticulations, thicke and moe ounded shafts, a lowe neck-shaft angle and a boade poximal shaft than in moden humans (Ruff and Walke, 1993; Ruff et al., 1993; Weave, 2002, 2003). Fom the publications on ace assessment discussed above, a clea elationship has not been demonstated between femoal cuvatue and climate (Bookstein et al., 2003) but it is woth consideing this again in light of the cuent eseach, though investigating the possible elationship between oveall skeletal mophology and long bone cuvatue. Long tem climatic adaptation may have an impotant effect on the size and shape of long bone diaphysis (eason, 2000b). 34

66 2.4. Hypotheses and pedictions The backgound povided by the two peceding Chaptes (1 and 2) on bone cuvatue in Neandethals and moden humans suggests thee main hypotheses to explain vaiation in long bone cuvatue in ecent moden humans. These hypotheses and the associated pedictions will be the basis fo the analysis and ae listed below. Hypothesis 1: A high degee of cuvatue is elated to body size. Body size affects the mechanical loadings of weight-beaing skeletal elements and cosssectional diaphyseal popeties. Biewene (1983) suggested cuvatue is a mechanism by which lage animals educe bone stesses because body mass inceases moe apidly than the cosssectional aea of bones. Although this elationship is clea fo weight-beaing bones such as the femu, Swatz (1990) demonstated that cuvatue of the adius in anthopoids was also allometically elated to body size, and could not find a elationship between cuvatue and diffeences (tension o compession) in loading egime between bachiatos and non-bachiatos. Associated pedictions: - Body size is positively coelated with degee of femoal and adial cuvatue. - Males have highe degees of cuvatue than females, because males ae, on aveage, lage. Hypothesis 2: Cuvatue is a esponse to inceased activity levels Seveal pedictions follow fom the expected elationship of habitual behaviou of long bone cuvatue. Males have highe activity levels than females, especially in hunte-gathee societies whee division of labou is most ponounced, and this may esult in sexual dimophism in cuvatue (Lasen, 1995). Activity levels in adults decease with age (Caspesen et al., 2000; Noman et al., 2002), so cuvatue may also decease with inceasing age. 35

67 Habitual use of the foeam in a pat-flexed position duing ponation, with the hand in supination o semi-supination, esults in a moe medially placed adial tubeosity, inceased stain in the foeam and may esult in a elatively longe adial neck (Tinkaus, 1988). This inceased stain is expected to esult in moe cuvatue. Associated pedictions: - Males, having highe activity levels than females, also have highe degees of cuvatue. - Thee will be a positive coelation between cuvatue and obusticity. - opulations with highe levels of aquatic mobility will have the most lateally cuved adii and most posteioly cuved ulnae. - With inceasing individual age and deceasing activity levels, thee will be a decease in cuvatue. - With inceasing sedentism though time in Euope, thee will be a decease in cuvatue. - osition of the adial tubeosity and adial neck length will be coelated with cuvatue. - A highe degee of femoal cuvatue will be associated with a moe distal apex of cuvatue Hypothesis 3: Cuvatue is a consequence of adaptation to cold climate. Individuals in high latitudes have elatively shote distal limbs and elatively lage aticulations than those living in wam climates (Ruff, 1994b). The shape of the femu has been suggested to be a consequence of long tem climatic adaptations in the pelvis. The wide pelvis in coldadapted populations esults in elatively lage aticulations, geate shaft obusticity and low neck-shaft angles, as well as longe elative neck length and inceased tosion (Weave, 2003). Little is known about how cold adaptation affects the lowe am. Associated pedictions: - Thee will be a positive coelation between cuvatue and latitude (used as a quantitative measue fo aveage tempeatue). - Thee will be a positive coelation between cuvatue and obusticity of the epiphyses and shaft. - Thee will be a positive coelation between femoal cuvatue, elative neck-length and tosion and neck-shaft angle. 36

68 CHATER 3. MATERIALS AND METHODS 3.1. Mateials The mateials included in this study can be divided into two goups: 1) Neandethals and ealy anatomically moden humans, and 2) the compaative ecent moden human sample. The ecent moden human sample is a geogaphically and behavioually divese sample that was chosen to investigate the influence of climatic, body size/body popotions and activity levels on cuvatue Neandethal and ealy anatomically moden human fossils Neandethals and ealy anatomically moden human emains ae elatively abundant compaed to othe hominin fossils but the sample is smalle than would be ideal fo a compehensive compaative analysis. All available femoa, ulnae and adii wee studied, and whee the oiginal was missing o damaged, casts wee used. The sample is compised of complete o nealy complete bones Neandethals The sample of Neandethals epesents Middle alaeolithic Westen Euopean (so-called classic Neandethals) and westen Asian Neandethal sites (Table 3-1) dating fom 65Ka-35Ka B. A shot desciption and some key efeences fo each site is included below with the most ecent fist. 37

69 Table 3-1 Summay of the Neandethal sample, by egion. NEANDERTHAL complete specimens Adult Femu Ulna Radius Euope Spy 1 a X Spy 2 a X La Feassie 1 b X X La Feassie 2 b X X X La Quina H5 b X X La Chapelle aux Saints b X X X Le Régoudou c X X Levant Kebaa d X X N Adult cast Femu Ulna Radius Euope Le Moustie e X X X Neandethal f X X * X westen Asia Shanida 1 g X X Shanida 5 g X Shanida 6 g X N a Royal Belgian Intitute fo Natual Sciences, Bussels b Musee de l Homme, ais c Musee du eigod, éigeux, d Tel Aviv Univesity e Museum fü Vound Fühgeschichte in Belin f Rheinisches Museum in Bonn g Smithsonian Institute Washington, * pathological 38

70 Spy Two patial skeletons and some juvenile fagments wee discoveed in Spy, 15 km west of Namu, Belgium, in 1886 by M. Lohest and M. De uydt (Faipont and Lohest, 1887). The fossils wee associated with Mousteian tools (Bodes, 1959), but because of the ealy date of the excavation and poo excavation techniques, dating is poblematic. The fossils ae tentatively dated to Ka B based on associated faunal emains (Cody, 1988). Spy 1 is believed to be an adult male of appoximately 35 yeas old. The calotte, a patial maxilla and patial postcanial emains ae peseved. Spy 2, also a patial male skeleton, consists of a calotte and some isolated teeth and postcanial emains. Thee is some confusion about the postcanial elements and thei association with eithe Spy 1 o Spy 2. Only Spy 1 has a completely peseved adius and was included in the analyses. The othe specimens ae too fagmentay to be included. Both specimens ae undoubtedly Neandethals (Faipont and Lohest, 1887; Boule and Vallois, 1952). The Spy emains eside in the Royal Belgian Institute of Natual Sciences in Bussels, by coutesy of the family of ofesso Max Lohest ( ). La Feassie The site of La Feassie, Fance, was discoveed in 1909 by D. eyony and L. Capitan and yielded the emains of two adults (La Feassie 1 and 2) and possibly 6 o 7 juveniles (La Feassie 4a, 4b, 5: neonates o fetuses; La Feassie 3 and 7, possibly same individual: +/- 10 yeas old; La Feassie 6: +/- 3 yeas old; La Feassie 8: +/- 2 yeas old) (Heim, 1968). The emains wee found in a ock shelte 3.5 km fom Bugue, Fance, and wee associated with Mousteian tools and a cold-climate fauna. The site dates to appoximately 40 Ka B (Heim, 1968; uech, 1981) and the skeletal mateial could have possibly been intentially buied (eyony, 1934 in Schwatz and Tattesall, 2002). 39

71 La Feassie 1 is a patial skeleton of an adult male (+/- 45 yeas old) and La Feassie 2 is an adult female (25-30 yeas old) (Heim, 1968). La Feassie 1 is the best peseved but the femoa wee too incomplete to be included in the sample. The ulna and adius fom both the left and the ight side wee included. Fo La Feassie 2 the femoa, and adius and ulna fom the ight side ae included in the sample. Le Moustie The site of Le Moustie comes fom the village of Le Moustie, Fance, which is located about 10 km fom Les Eyzies de Tayac. The hominin emains wee discoveed by O. Hause in 1908 who late sold them to the Museum fü Vo- und Fühgeschichte in Belin. The ock shelte contained atefacts of the Mousteian tadition and is dated to /-2.6 Ka B using TL dating on bunt flint (Bodes, 1959; Valladas et al., 1986) and ESR dating on associated mammal bones (Mellas and Gun, 1991). Cut-maks and bone modifications indicate that Le Moustie 1 was killed intentionally by peimotal impacts, the head was decapitated, the mandible focibly disaticulated and the copse (obviously completely dismembeed) defleshed. (Ullich, 2005 p. 304). The adolescent skull is cetain to belong to a Neandethal. It has a low foehead, double ached bowidge, a low vault, lambdoidal flattening, a supainiac depession and an occipital bun. Duing WWII most of Le Moustie 1 was destoyed and only the skull and some of the postcanial elements emain (Day, 1986; Schwatz and Tattesall, 2002). Due to a watime fie the oiginal fossils ae heavily distoted. Reliable measuements can only be taken on casts made fom the oiginals (Thompson and Nelson, 2000). Only a plaste cast of the econstuction of the left femu, ight ulna and ight adius ae complete enough to be included in the sample. Thomson and Nelson (2005) emaked on the exaggeated length of the cast of the adius and the econstuction of both extemities. They descibe the adial shaft as stongly lateally cuved and having a medially oiented adial tubeosity. The oiginal adius was missing most of the epiphyses but they have been econstucted on the cast. The ulna is mostly peseved. The tochlea notch faces anteioly. Thee is no clea adial notch on the cast. The femu is patly econstucted. The lesse tochante, geate tochante, 1/3 of the femoal head and most of the distal epiphyses ae econstucted. 40

72 Neandethal (also Feldhofe) The Feldhofe Neandethal is the type specimens fo Homo neandethalensis and was found in 1856 by wokmen fom a quay in the Neandethal Valley, about 11 km east of Düsseldof, Gemany. Neithe atifacts no mammalian bones wee ecoveed fom the site, although eexcavation of the old mining deposits since 1998 (Schmitz et al., 2002; Schmitz, 2006) evealed stone tools and faunal emains along with moe Neandethal emains (Day, 1986; Schmitz et al., 2002). Thee ae now thee individuals epesented fom the site. On the basis of mtdna analysis of the oiginal Feldhofe emains Kings and colleagues (1997) demonstated that the Neandethal genome was diffeent fom that of moden humans. Futhe mtdna analyses of the moe ecently discoveed Feldhofe emains yield sequences simila to those of othe Neandethals and ae diffeent fom those of moden humans (Schmitz, 2006). Cabon-14 dating of the newly discoveed emains indicates an age of appoximately 40 Ka B. The Feldhofe 1 skull has a clea Neandethal anatomy, and sutue fusion suggests an age of appoximately 50 yeas at death (Day, 1986; Schwatz and Tattesall, 2002). The postcanium includes two femoa, two ulnae and the ight adius. Although thee is evidence of slight defomation on the femoa and the adius, the left ulna is too pathological to be included in the study. The long bones ae thick and show ponounced muscula attachments. The humei ae staight but the adius is cuved and has a lage adial tubeosity. The factue elated pathology on the left elbow would have limited the movement of the joint. The femu is cylindical and shows signs of a thid tochante (Heim, 1981, 1982, 1983). La Quina H5 The site of La Quina, 25 km south of Angoulème, Fance, was found in 1872, but it was not until 1908 that Heni-Matin discoveed the fist hominid emains (Matin, 1921). A total of 27 individuals ae peseved; howeve, only one individual, H5, is included hee (left ulna and adius). H5 is a patial adult skeleton that was found associated with Mousteian of the La Quina tadition (Debénath et al., 1998). Although the hominin emains come fom diffeent layes, H5 41

73 comes fom the ealie levels belonging to OIS 4 and dating to appoximately 65 Ka B (Mellas, 1996). A moe ecent date of Ka B based on chonometic data has also been epoted (Zilhao, 2006). Shanida In 1951 R. Solecki discoveed the site of Shanida in the Zagos Mountains in Iaq, appoximately 400 km noth of Baghdad. The emains of at least nine patial skeletons wee found in a lage cave (Solecki, 1957, 1961, 1975). Although moden human buials wee discoveed in the uppe layes of the site, the Shanida Neandethals wee found in a single laye associated with Mousteian tools, heaths and local fauna (Solecki, 1957, 1961). The Mousteian Neandethal laye was 14 C dated to appoximately 50.6 Ka B (Ba-Yosef, 1989). Six adults, one young adult and two infants wee found at the site and wee descibed by Stewat (Stewat, 1962, 1963, 1977) and Tinkaus (Tinkaus, 1978, 1982a, 1982c, 1982b, 1983b). The skulls show a long low canial vault, a lage supaobital tous, mid-facial pognathism, a tansvese occipital tous and a ounded vault in occipital view. The mandible lacks a chin and the anteio teeth ae heavily won. Because of these featues, thei classification as Neandethals has neve been questioned (Solecki and Solecki, 1974; Solecki, 1975; Stewat, 1977; Tinkaus, 1978; Tinkaus and Zimmeman, 1979; Stinge and Tinkaus, 1980; Tinkaus, 1982a, 1982c, 1982b; Tinkaus and Zimmeman, 1982; Ivanhoe and Tinkaus, 1983; Tinkaus, 1983a). The post-cania fom the site show a high degee of obusticity and display signs of poweful musculatue. The sample used hee includes the left ulna and adius of Shanida 1, the ight ulna of Shanida 5, and the left adius of Shanida 6. Because of the cuent elocation of the mateial fom the Baghdad Museum, casts of this mateial wee measued at the Smithsonian Institution. La Chapelle-aux-Saints This patial Neandethal skeleton was discoveed in 1908 by A. and J. Bouyssonie and L. Badon nea the village of La Chapelle-aux-Saints, 40 km fom Bive, Fance. It was found buied in a cave (Badon et al., 1908 in Schwatz and Tattesall, 2002) and associated with an 42

74 advanced Mousteian industy and mammals epesentative of a tempeate climate (Boule, 1908). The laye fom which the specimen came has an absolute date of Ka B using ESR dating on mammal teeth (Gün and Stinge, 1991). The skeleton is faily complete and belonged to an aged adult male. The specimen has typical Neandethal featues such as an occipital bun, supa-iniac fossa, small mastoid pocesses and mid-facial pojection (Boule, 1908; Tinkaus, 1985). The ight side of the postcanial skeleton is well peseved, and the ight femu, ulna and adius ae included in the sample. Thee ae signs of degeneative joint disease in the skeleton consistent with its infeed old age (Tinkaus, 1985). In geneal, the long bones ae shot and thick with stong muscle makings and shot distal limb segments compaed to its poximal limb segments. The humei ae staight but the femu and adius ae bowed (Tinkaus, 1985). Kebaa The Mughaet el-kebaa is appoximately 13 km south of Wadi el-mughaa on the westen slope of Mount Camel in Isael. The excavation of the site began in Duing the ealy stages of the excavation the fagmentay emains of an infant wee discoveed (Kebaa 1). In 1983, an adult Neandethal buial was ecoveed (Kebaa 2 commonly efeed to as simply Kebaa) (Goldbeg and Ba-Yosef, 1998). Although the skull and most of the lowe limbs ae missing the skeleton is well peseved. The skeleton is estimated to be that of a yea old male individual. The pelvis indicates that Neandethal pelves ae fundamentally diffeent fom moden human ones, even when compaed to moden humans fom the same time peiod. They have a long supeio pubic amus which pobably stems fom a moe extenally otated innominate bone and may be attibuted to diffeences in locomotion and postue elated biomechanics (Rak and Aensbug, 1987; Rak, 1990). The laye fom which the adult buial oiginates dates to appoximately Ka B (Goldbeg and Ba-Yosef, 1998). The occupation laye also contained Mousteian tool technology (Ba-Yosef et al., 1986). The adius and ulna fom both sides ae included in the sample. The patially peseved femu lacks its distal epiphyses and is too fagmentay to be used. 43

75 Le Régoudou In 1957, R. Constant discoveed a collapsed limestone cave 2 km noth of Montignac, Fance, containing Mousteian tools, and the emains of two individuals: one patial skeleton of a young adult (Régoudou 1) and some pedal elements (Régoudou 2) (iveteau, 1959). The site was eexcavated by Bonifay fom 1957 onwads and based on the sedimentology, fauna and Middle alaeolithic technology he assigned the specimen to OIS 4 (oughly 65Ka B) (Bonifay and Vandemeesch, 1962; Bonifay, 1964). The individual is pobably a young adult in its mid-twenties. It is not possible to detemine its sex as the canium and the pelvis ae pooly peseved. The ight ulna and adius ae complete enough to include in the sample. The canial mophology shows a clea Neandethal affinity as does the mophology of the postcanial skeleton (iveteau, 1959; Vandemeesch and Tinkaus, 1995). 44

76 Ealy moden humans The sample epesents ealy anatomically moden humans fom Euope and westen Asia (Table 3-2). A shot desciption and some key efeences fo each site is included below in chonological ode fom most ecent to oldest. Table 3-2 Summay of Ealy Moden Human sample, by egion. complete specimens Adult Femu Ulna Radius Euope Abi ataud a X X Chancelade b X X Combe Capelle b X X X Westen Asia Sungi c X X X avlov d X Dolni Vestonice 13 d X X X Dolni Vestonice 14 d X X Dolni Vestonice 15 d X* X* X* Dolni Vestonice 16 d X X X Levant Ein Gev e X X Ein Gev Nahal e X Ohalo II e X X X Qafzeh 9 e X X X Skhul IV f X X N Adult cast Femu Ulna Radius Euope St. Gemain g X X X Westen Asia Kostienki 14 h X X X N a Musee de l Homme, b Musee du eigeux, c Laboatoy fo econstuction, Moscow d Dolni Vestonice, e Tel Aviv Univesity, f Havad eabody Museum, Boston, USA, g Musée National du ehistoie, h Kunstcamea St etesbug *pathological 45

77 Ein Gev The site of Ein Gev is 1 km east of the Sea of Galilee in nothen Isael. The site was excavated oiginally by Stekelis and Ba Yosef in The achaeology at the site is Epipalaeolithic Kebaan, and 14 C dating on chaed bone indicates an age of B +/-415 (; Davis, 1974). The human emains at the site come fom a buial and pobably belonged to an adult female (30-40 yeas old) (Stekelis and Ba-Yosef, 1965). The bones wee quite fagmentay at the time of discovey, but most pats could be estoed. Despite the estoation it was only possible to include the ulna in the analyses (Aensbug and Ba-Yosef, 1973). Chancelade In 1888 M. Hady discoveed a Magdalenian skeleton at the site of Raymunden in the village Chancelade, nea éigeux, Fance (Sollas, 1927; Billy, 1969). The deposits ae believed to be a buial and the skeleton is epoted to have been coveed with oche. The almost complete skeleton is that of a yea old man who was appoximately 1.6m tall. The canium was once mistakenly believed to be that of an Eskimo and the Eskimo-like featues wee intepeted in light of the cold envionment duing the Magdalenian Age (Testut, 1925 in Keith, 1925; Sollas, 1927). The skull is clealy that of a moden human and is associated with an achaeological deposit of Magdalenian III o IV, dating most pobably between Ka B (Ruff and Walke, 1993; Tinkaus et al., 1999a). The associated fauna ae indicative of cold conditions but an absolute date fo the site has not yet been established. The postcanial emains wee descibed by Billy (1969). Subsequent publications by othe authos have demonstated some of the highest values fo obusticity found in any ealy moden human (e.g. Ruff and Walke, 1993; Tinkaus et al., 1999a). The left femu was pooly econstucted and extemely fagile but the ight femu is included in the sample as well as the ight ulna, although thee was some econstuction of the femoal head and distal condyles but none of the landmaks wee affected. 46

78 Saint-Gemain-la-Rivièe The site of Saint-Gemain-la-Rivièe, Fance, is an ealy Magdalenien ock shelte and dated to between Ka B using 14 C dating (Costamagno, 2002; Ducke and Heny-Gambie, 2005). It was excavated on and off between 1929 and A complete adult human skeleton of was discoveed in 1934 (Blanchad, 1935; Vaufey, 1935). The skeleton was discoveed in a buial stuctue made out of ocks and was adoned with maine shells and teeth of ed dee and eindee. The skeleton is believed to be that of a young adult female (Vaufey, 1935; Heny- Gambie et al., 2002). Cabon and nitogen isotope atios in the bone collagen of the young woman indicate that the main souce of potein was lage hebivoes. She did not consume significant amount of fish and he subsistence patten eflects a less oppotunistic diet that geneally attibuted to humans fom the ealy Magdalenian (Ducke and Heny-Gambie, 2005). The oiginal fossils wee not available fo study, so a cast was measued. The left ulna, ight adius and ight femu wee included in the sample. The patella of the ight femu is fused to condyles but did not affect the landmak collection. Ein Gev Nahal Nahal Ein Gev is an Uppe alaeolithic buial of an almost complete skeleton in the noth of Isael. The associated achaeology is Levantine Auignacian, which places the individual in the Uppe alaeolithic athe than Epi-alaeolithic. Diect dating of the emains has not been successful but sites with simila deposits, such as Ohalo II, have been dated to 19Ka B (Aensbug, 1977). The skeleton is believed to be that of a yea old female. She had gacile canial featues and shot statue (appox. 157 cm). The skull is diffeent fom othe Uppe alaeolithic cania in its size and shape. Mophologically, it is most simila to Co-Magnon II and edmostí IV, which has been suggested to be an indication of common ancesty (Aensbug, 1977; Belfe- Cohen et al., 2004). The emains ae badly damaged and most of the long bone epiphyses wee cushed. Because of this extensive damage, only the ight adius was sufficiently econstucted to be included in the sample. 47

79 Abi ataud 6 The ock shelte of Abi ataud was found in the town of Les Eyzies de Tayac in 1958 by H. Movius (Movius, 1966, 1975). Thiteen individuals wee ecoveed, howeve, most of these ae incomplete. The best peseved specimens ae a female canium, ataud 1, and an adult skeleton, ataud 6. The most ecent estimated date fo the site is between Ka B (Movius, 1966, 1975; Mellas et al., 1987; ettitt et al., 2003). The human emains most pobably come fom the uppe levels of the site and, if this is coect, would date to appoximately 22 Ka B (Mellas et al., 1987; ettitt et al., 2003). The emains wee associated with a oto-magdelenian industy (Movius, 1966). In the cuent study only the left ulna and adius of ataud 6 wee used. Ohalo II Ohalo is an Uppe alaeolithic site in the Levant nea the Sea of Galilee that dates to 23,500-22,500 B base on adiocabon dating (Nadel and Heshkovitz, 1991). Excavations evealed bush huts, heaths and a human gave. Ohallo II H2 is a elatively complete adult male skeleton estimated to have been between 35 and 40 yeas at death. The left adius and ulna wee damaged and only the ight side is included in the sample (Heshkovitz et al., 1995). Sungi (also Sounghi) The Sungi site has been excavated since 1957 and is located appoximately 200 km notheast of Moscow. It has yielded both a single and a double buial. The single buial is that of an adult male (Sungi 1). The double buial is that of an adolescent male and female (Sungi 2 and Sungi 3, espectively). All thee buials buials wee in extended, supine position. Sungi 2 and 3 wee lying head to head and wee coveed in ed oche. The buials have been diectly dated using adiocabon dating. Sungi 1 is Ka old, wheeas the Sungi 2 and 3 double buial is Ka old and thus slightly olde than Sungi 1 (ettitt and Bade, 2000; Ovchinnikov and Goodwin, 2003 but see Kuzmin et al., 2004). The 48

80 Sungi 3 gil has a pathology that is emakably simila to the obseved chondodysplasia calcificans punctata of Dolni Vestonice 15 (pesonal obsevation, Tinkaus et al., 2001). The pathology pesents itself with sevee skeletal defomaties of the long bones. The ight femu, ulna and adius of Sungi 1, the adult male, is included in the sample. avlov I The site of avlov, containing two skeletons (avlov I and avlov II), is located close to Dolni Vestonice and appoximately 35 km South of Bno, Czech Republic. The site was found and excavated in 1952 by B. Klima. The tool industy at the site is known as Easten Gavettian (Vlcek, 1961a, 1961b, 1991; Svoboda, 1994; Adovasio et al., 1996). The alov I skeleton is an adult, most likely male, and includes a patial canium, maxilla, mandible, isolated teeth and a patial skeleton. The buial dates to Ka B based on adiocabon dating (Klima, 1987). The emains ae believed to be those of an ealy moden human. Because it is a faily obust skeleton and canial featues, such as oveall obusticity and a swollen sub-lambdoidal aea eminiscent of an occipital bun, the avlov skeleton has been suggested to be a link between achaic Euopeans (Neandethals) and moden humans (Smith et al., 1982; Wolpoff, 1996). Only the ight adius of avlov I was sufficiently well peseved to be included in the sample. Dolni Vestonice (also Dolni Vĕstonice) Dolni Vestonice is a complex of sites in and aound the village of Dolni Vestonice, 35 km South of Bno in the Czech Replublic. The sites wee discoveed by Absolon in 1925 and late excavated by Klima fom Thee ae 16 individuals epesented at the cluste of settlements and they pobably all date to appoximately 26.5 Ka B (Svoboda and Vlcek, 1991; Fomicola et al., 2001). Thee ae two aeas at the site: one containing most of the occupational infomation and one with the human emains. The associated industy is Gavettian, which is accompanied by engaved bone tools and clay figuines. Most of the human emains ae buials. The tiple buial of 49

81 individuals XIII, XIV and XV is that of thee young, possibly genetically elated, adults who wee buied togethe with gave goods. The cental skeleton, which is pobably a female, has pathologies causing sevee bone defomation of the femoa and foeam (chondodysplasia calcificans punctata, Tinkaus et al., 2001). The othe individuals ae males (Klima, 1987; Bahn, 1988; Alt et al., 1997; Tinkaus et al., 2000; Fomicola et al., 2001). Although the human emains ae consideed to be moden humans and ae elatively gacile, some authos have suggested that they etain pimitive and Neandethal-like featues and ae indicative of continuity in the egion (Smith et al., 1982) All thee individuals fom the tiple buial ae elatively well peseved and have at least one well peseved femu, ulna and adius. Because of its sevee pathology, Dolni Vestonice XV was excluded fom the analyses. The left femu, ulna and adius of Dolni Vestonice XVI, both femoa, ulnae and adii of Dolni Vestonice XIII, and both femoa and ulnae and the left adius of Dolni Vestonice XIV wee included in the sample. The sepaate femoal head of Dolni Vestonice XIV was held in place duing data collection. Combe Capelle The ock shelte of Combe Capelle, 20 km Southeast of Begeac, Fance, was discoveed in It has yielded a patial hominin skeleton dated to appoximately Ka yeas (Valladas et al., 2003). The skeleton was associated with Gavettian tools and its mophological affinities ae clealy moden (Lenoi and Dibble, 1995). The skeleton was lost duing the same fie that destoyed the Le Moustie adolescent Neandethal emains, but in 2002 the skull was ediscoveed in the museum. Regetfully, the postcanial skeleton is still missing (Hoffmann and Wegne, 2002). Howeve, thee ae well peseved oiginal plaste casts of the left and ight femoa and ulnae that wee included in these analyses. Thee ae mino aeas of the oiginal bone that ae damaged, such as some abasion of the distal femoal condyles and the lack of the styloid pocess, but this should not seiously affect the esults. The ight adius was also complete enough to be included. 50

82 Kostenki 14 (also Makina Goa) The site of Kostenki (Makina Goa) is in the Voonezh egion in Russia. The site is pat of a complex of sites that povides an impotant statigaphic sequence fo the egion between the Capathian and Ual Mountains (Sinitsyn and Hoffecke, 2006). It has yielded a numbe of skeletal emains: a 5-6 yea old child (Kostenki 15), an eldely man (Kostenki 2), a 9-10 yea old child (Kostenki 12) and a well peseved skeleton of a young adult male (Kostenki 14). Kostenki 14 was discoveed in a gave and coveed with yellow and ed oche (Jelinek et al., 1969). The skeleton came fom the lowemost cultual layes at Makina Goa and is adiocabon dated to at least Ka B (Sinitsyn, 2003). It is a young male that was pobably aound 160 cm tall. The supaobital tous is moden-human-like. Jelinek (1969) also descibes Kostenki 14 as being simila to the emains fom Gimaldi and Co-Magnon. Although the emains ae not cuently available fo eseach because a monogaph is in pepaation, the cuato at the Kunstkamea in Saint-etesbug, Russia, allowed the inclusion of the casts in the analyses. The ight femu and ulna and the left adius ae sufficiently peseved to be analysed. Qafzeh (also Jebel Qafzeh) The site of Qafzeh in Isael was discoveed in 1933 by R. Neuville (Vandemeesch, 1981). The site is 2.5 km south of Nazaeth and is located on Mount Camel. Up to 12 individuals have been discoveed in the cave. The tool industy is Levalloiso-Mousteian with some backed knives and buins of Uppe alaeolithic chaacte (Vandemeesch, 1981). Thee is a themoluminescence date of 100 Ka B +/- 10 Ka (Gün and Stinge, 1991). The human emains belong to eight adults (Qafzeh 1,2,3,5,6,7,8,9), thee infants (Qafzeh 4, 4a, 10) and one ten-yea old child (Qafzeh 11). A detailed desciption of the human emains can be found in Vandemeesch (1981). In geneal, the postcanial featues ae moden and do not show distinct Neandethal o othe achaic featues (Vandemeesch, 1981). Tinkaus suggests, howeve, that both Qafzeh and Skhul (see below) have a mosaic of featues and agues that 51

83 mophological and achaeological evidence can best be explained by continuity between achaic and moden humans (Tinkaus, 1981). The postcanial emains of Qafzeh 9, an adult male, wee complete enough fo the ight femu and ulna, and left adius to be included in the analyses. Skhul (also es-skhul) At least 10 individuals wee found at the site of Mughaet es-skhul (usually efeed to in the liteatue as Skhul ) on Mount Camel, in southeasten Isael. The site was discoveed in 1929 duing an excavation diected by D. A. E. Gaod (Gaod and Bate, 1937). Most of the bones wee associated and showed little distubance, indicating that they wee pobably buied intentionally (Gaod and Bate, 1937) The emains ae associated with the Levalloiso-Mousteian and the fauna is simila to that at the adjacent site of Tabun. Mean ESR age estimates place the site between 81 and 101 Ka B (Gün and Stinge, 1991) and TL dating dates the site to an aveage of 119 Ka B (Valladas et al., 1998). At the site, seven adults and thee juvenile individuals ae epesented. Most of these ae patial skeletons and ae consideed to be anatomically moden. The skeletons ae long and slende compaed to Neandethals. Thee ae some pimitive featues pesent, though, such as the stout foot and finge bones and well-developed thumbs (Vandemeesch, 1981; Tinkaus, 1993; Niewoehne, 2001). One theoy attibutes the appaent pesistence of these to inbeeding between ealy modens moving into the egion fom Afica, and Neandethals coming in fom Euope (Kame et al., 2001). Altenative views see the Skhul and Qafzeh people as membes of an ealy moden population that evolved in the Levant (Vandemeesch, 1981; Rightmie, 1998). The adius and ulna of Skhul IV ae included in the sample. 52

84 Moden populations The moden human compaative sample was chosen specifically to test hypotheses of factos that explain long bone cuvatue (see Chapte 2 fo details). The sample consists of adult femoa, ulnae and adii. All the individuals in this sample ae skeletally adult based on closue of the epiphyses and pathological individuals wee excluded. Fo 93 individuals the age at death was known. Sex of the individuals was ecoded fom the museum catalogues o, if pelvis and canium wee available, sex was detemined by obsevation. Individuals whee sex detemination was impossible and whee the museum had no infomation wee labelled as unknown. The elatively small numbe of individuals pe population is due to the availability of postcanial mateial in museum collections. In ode to captue the ange of vaiation in moden humans thoughout the wold, some small samples wee included as pat of goups ceated fo futhe analyses (See section 3.2 in this Chapte). Whee possible the femu, ulna and adius fom the same side of the skeleton wee included in the sample. When this was not possible bones fom opposite sides of the skeleton wee included. Table 3-3 below eflects the total numbe of individuals epesented in the sample athe than the numbe of bones. Sample numbes of paticula bones ae specified in the esults chaptes. 53

85 Table 3-3 Summay of ecent moden human sample, alphabetically. opulation N Collection Location Afican-Ameican 12 Afican-Ameicans Tey Collection Smithsonian, Washington Alaskan Aleut 15 Aleutian Islands Collection eabody, Havad Andaman Islands 17 College of Sugeons Collection NHM, London Aizona Native 20 Canyon del Muetos NHM, New Yok Austalian 13 College of Sugeons Collection NHM, London Aboiginals Belgian Medieval 29 Spy and Gutschoven RBINS, Bussels Belgian Neolithic 72 Fufooz, Mauenne, Hastièe, RBINS, Bussels Dinant Bitish Neolithic 2 Coldum NHM, London Chinese 9 Chinese Cemetay, Kaluk Quad Alaska Smithsonian, Washington Coloado Native 4 Montezuma County, Coloado eabody, Havad Czech Medieval 39 Moavian Empie Collection NHM, ague Danish Medieval 15 Sankt Bendtskike, Ringsted Univesity, Copenhagen Danish Neolithic 49 Koshoj Adby, Guldhoj, Boeby Univesity Copenhagen Egyptian 5 Egyptian Dynasty NHM, ais English Medieval 21 Scaboough NHM, London English Uban 21 Spitalfields 18th-19thC NHM, London Fench Medieval 16 Villeboug, St. Gabiel NHM, ais Fench Neolithic 24 Valée du etit Moin NHM, ais Geenland Inuit 31 Tuqutut, Ilutalik, Uunatoq, Ilosuit Univesity, Copenhagen Khoi o KhoiKhoi 10 Oxfod Collection NHM, London Kazach 7 Southen Volga Region St. etesbug Lapland 17 Russian Saami Moscow State Univ. Natufian 16 Mallaha Univesity, Tel Aviv New Mexico 9 Aztec Ruins NHM, New Yok Ohio Native 18 Madissonville, Ohio eabody, Havad euvian 13 Ancon (Lima) NHM, ais oint Hope Alaska 15 Alaskan Inuit NHM, New Yok ygmy 4 Litui Cental Afica RBINS, Bussels Russian Eskimo 15 Sibeian eninsula, Ekveni Moscow State Univesity Russian Mesolithic 22 Vasilievski St.-etesbug Sibeia 16 Sibstey, Salehad Sibeia Moscow State Univesity South Dakota Native 13 Campbell County, Ohae Resevoi Smithsonian, Washington Tasmanian 2 Tasmania NHM, London, Bussels Tiea del Fuego 2 Tiea del Fuego, Agentina NHM, Vienna TOTAL 593 individuals NHM = Natual Histoy Museum; RBINS= Royal Belgian Institute of Natual Sciences 54

86 Afican-Ameican The Afican-Ameican sample is fom the Tey collection. It was collected by Robet J. Tey ( ) fom a local St. Louis hospital and institutional mogues. The mateial in the collection consists pimaily of uban living individuals whose bodies became popety of the state when they wee not claimed, o whose elatives signed ove the bodies to the state. The Tey collection consists of 1728 individuals of known age, sex, ethnic oigin, cause of death and pathological conditions and twelve individuals wee andomly sampled. Alaskan Aleut and oint Hope Alaskan The Alaskan Aleut and the oint Hope Alaskan ae achaeological samples. The Alaskan Aleut ae membes of the Inupiak, a subdivision of the Inuit. They taditionally lived in goups of in the nothen actic egion and elied mainly on lage sea mammal hunting fo subsistence. They hunted these animals with stone, bone, ivoy and wooden tools such as hapoons, aows and knives. Thei diet is almost entiely canivoous, as thee is vey little plant mateial available in the aea. Some populations have been found to eat sea weeds and gasses o the stomach contents of the animals hunted (Buch and Buch J., 2006). Andaman Islands The Andaman Islands ae located in the Indian Teitoial pat of the Bay of Bengal. The Andamanese ae hunte-gathees, who ely on eating indigenous mammals, plants and fish acquied with stone, bone, wooden tools and nets (Radcliffe-Bown, 1948). Aizona Native Ameicans The Native Ameindians fom Aizona come fom a site called Canyon del Muetos, Tempe. The Los Muetos site was occupied by the Hohokom cultues and dates to appoximately 500AD 1500AD (Hauy, 1945). Analysis of the palaeo-envionment of cental Aizona suggests that as 55

87 ealy as 750 AD the climate was aid and the iigation canals dating back to 1150AD indicate that hoticultue faming may have been pacticed at the site (Hauy, 1945). Austalian Aboiginals The Austalian Aboiginal emains ae cuated at the Natual Histoy Museum in London. The individuals come fom a vaiety of places in Austalia and ae pe- and post-contact. Although the individuals may have had diffeent cultual backgounds, most tibes wee teestial foages and will be teated in the analyses as such. They used speas and thowing sticks to acquie thei foods and lived in semi-nomadic villages (Jupp, 2001). Belgian Medieval The Belgian Medieval sample is cuated at the Royal Belgian Institute of Natual Sciences in Bussels. The sample comes fom two vey small ual villages in Belgium: Spy Bastin (13 th C AD) and Gutschoven (Caolingian Empie AD). They wee all fames o caftsmen (pesonal communication, Semal). Belgian Neolithic The Belgian Neolithic sample is cuated at the Royal Belgian Institute of Natual Sciences in Bussels. The Belgian Neolithic sample compises individuals fom the Middle and Late Neolithic peiod (+/ B to +/ B cal. in the Seine-Oise-Mane distict). The specimens come fom fou diffeent sites with Dinant being the oldest ( B) and Fufooz being the youngest ( B) (Cauwe et al., 2001). Although the sample comes fom gaves in ock sheltes o in the open ai and fom settlements oganised aound flint mines, it is believed that all the individuals had simila agicultual lifestyles (Toussaint et al., 2001). 56

88 Bitish Neolithic The Bitish Neolithic sample was extemely fagmentay. The sample was collected fom a mass gave in Coldum, Kent, and its fagmentay natue is due to the emoval and ebuial of the emains duing ceemonies. The sample skeletal mophology suggests that duing this peiod in England thee was a shift to agicultue fom mixed foaging but pecise infomation on this population is not available (Clinch, 1904; Wysocki and Whittle, 2000). The sample dates back to appoximately BC (Whittle et al., 2007). Chinese This 20 th centuy Chinese sample was cuated at the Smithsonian Institution, Washington D. C.. The emains wee collected fom the Kulak Cemetey in Alaska and consists only of males. This Chinese cemetey was used to buy the emains of the Chinese laboues that woked at a local fish canney. These Chinese ae assumed to be shot-tem immigants thee as thee is no sign of females o childen in the cemetey. Most of the settles came fom Cantonese Southen China (Hdlicka, 1944). Coloado Native This sample was cuated at the eabody Museum, Havad Univesity, Boston. The Native Ameindians fom Coloado come fom a site appoximately 15km fom Cotez in Montezuma County, south-westen Coloado. Although ethnic affiliation was not cetain, most of the county was inhabited by the Anasazi and the site dates back to Basketmake III ( AD). They wee mainly teestial hunte-gathees (Cum, 1996). Due to poo pesevation few individuals fom this population could be included in the sample. Czech Medieval The sample of Czech Medieval is cuated at the Natual Histoy Museum in ague. The individuals come fom the time of the Geat Moavian Empie (9 th C AD end 10 th C. AD) 57

89 (Dekan, 1981) and ae believed to come fom a faming population that lived on the lands suounding one of the bughs (pesonal communication,. Veliminsk, cuato). Danish Medieval This sample comes fom a cemetey in Denmak (Sankt Bendts Kike in Ringsted) and is cuated at the Medical Univesity of Copenhagen. The mateial was excavated in 2000 and dates back to 1080 AD ealy 1100s AD. At the time, faming was the main souce of subsistence, although it was fequently supplemented by the consumption of fish. The mateial has not been published (but see anum Baastup, 2002). Danish Neolithic The Danish Neolithic (appoximately 3000 BC) sample is a collection fom diffeent sites thoughout Denmak. The emains included in this poject ae fom Koshoj Adby, Uggeslev, Guldhoj and Boeby Island. The individuals lived in small settlements. Although fish was most impotant duing the Mesolithic, thee is evidence fo a dietay shift, and the Neolithic diet consisted mainly of teestial food which was hunted, famed and bed (cattle) (ia Bennike, pesonal communication; Böste et al., 1956; Taube, 1981; Richads et al., 2003). Egyptian The Egyptian sample date to the Old Kingdom and ae cuated at the Musée de l Homme in ais. The catalogue indicated that the individuals wee low status mummies fom the Old Kingdom (3000 BC). The Old Kingdom Egyptians wee intensive agicultualists gowing cops along the Nile Valley using iigation systems (Kamil, 1996). 58

90 English Medieval The Medieval Bitish sample comes fom Scaboough Castle Hill. It is a Medieval lay cemetey sample dating to the middle to late Medieval peiod (11 th -16 th Centuies AD). The lay individuals buied at the site pacticed faming and some fishing (Little, 1943; Mays, 1997). English Uban This is a sample of late 18 th ealy 19 th centuy Huguenots fom the cypt of Spitalfields chuch in London, England (Molleson and Cox, 1993). The individuals included in this sample ae all named adults with known ages at death. Individuals of both sexes and fom diffeent ages wee andomly selected. The population was an uban population of caftsmen and mechants. Fench Medieval The sample comes fom two sites, Villeboug in Cental Fance and St. Gabiel in the South of Fance and is dated to both the Meovingian ( ) and Caolingian peiod ( AD). Both samples ae assumed to have been fames although little is known about them (pesonal communication,. Mennecie, cuato). The sample has not been sexed o aged. Fench Neolithic The Fench Neolithic mateial comes fom multiple buial sites in the Valée du etit Moin, nothen Fance. The aea has a long agicultual histoy and these individuals ae believed to have pacticed intensive agicultue. The sample was collected duing the 19 th Centuy and elocated afte the Second Wold Wa fom the Musée des Antiquités in St. Gemain des és, Fance. The collection is substantial, but none of the postcanial bones ae individually catalogued no is thee any infomation available othe than the time-peiod (pesonal communication Mennecie, Bails). Theefoe, each bone is consideed as a sepaate individual. 59

91 Geenland Inuit The Geenland Inuit sample comes fom seveal coastal sites in Geenland: Tuqutut, Ilutalik, Uunatoq and Ilosuit. opulations fom these sites ae all pehistoic and had taditional Inuit lifestyles, elying mainly on fish and sea mammals fo thei subsistence (Bennike, 2006 pesonal communication). The postcanial emains ae not stoed individually, and little o no infomation is known on age o sex. Each bone is consideed a sepaate individual unless taphonomy and size made it possible to identify cetain sets of bones to belong to a single individual. Hottentot (also Khoi o Khoikhoi) This sample is cuated at the Natual Histoy Museum in London. The Khoi o Khoikhoi have been histoically efeed to as Hottentots. They ae a histoical division of the Khoi-San goup fom southwesten Afica. The Khoi wee pastoalists and pacticed animal husbandy of sheep, goats and cattle. This made it possible fo them to live in lage goups than suounding huntegathee populations. They gazed thei animals on the lage open plains until they wee foced into moe aid land by the expansion of the Bantu into Southen Afica (Boonzaie et al., 1996). Kazach The Kazach sample comes fom a pehistoic site in the Southen Volga ive egion in pesentday Kazachstan. Little is known about the collection othe than that the individuals most pobably led a taditional lifestyle of nomadic pastoalism (pesonal communication, J. Chistov). Lapland Saami (Also Sami o Lapps) This sample is fom the Kola eninsula and is believed to be pe-histoic (pesonal communication, 2007, D. ezhémsky). The Sami, also efeed to as Lapps, ae indigenous people of the Noth of Euope, and live in an aea coveing the noth of Sweden, Noway, Finland an the Kola eninsula in Russia. They wee taditionally nomadic and elied on a ange of subsistences: fishing, tapping, sheep and eindee heding, etc. 60

92 ( last accessed 18/01/2008). They ae a genetically distinct goup and wee pobably the fist to inhabit this nothen aea shotly afte the glacial ice eteated (Ingman and Gyllensten, 2007). Natufian The postcanial specimens fom the sites of Hayonim and Ein Mallaha ae extemely fagmentay so only a small sample could be collected. The Natufian is a Mesolithic cultue that existed in the Levant between Ka B. They ae thought to have built pemanent settlements befoe the onset of agicultue. This is evident at sites such as Hayonim and Ein Mallaha, whee living stuctues fom villages wee alongside buial stuctues. The Natufian wee teestial hunte-gathees and havested wild ceeals and gasses and tended to live close to pemanent wate souces. This havesting of wild ceeals is thought to eflect the onset of agicultue Muno (2004). The Natufian used stone tools that wee pedominantly micolith but also made sickle blades, ginding stones and bone tools such as hapoons and fish-hooks (Ba-Yosef, 1998; Muno, 2004). New Mexico Native Ameican The sample of pe-contact Native Ameicans fom New Mexico is a collection of an unidentified population, but the emains wee mistakingly associated with the Aztec Ruins (an Anasazi village misnamed Aztec see and have not yet been studied (Liste and Liste, 1990). The association to the Anasazi and the knowledge that ueblo also lived in the egion make it difficult to establish which cultual goup these individuals came fom. In any case, thee ae similaities in lifestyles between these goups: most peoples of this egion lived in pemanent o semi-pemanent settlements and wee agicultualists (G. Sawye, 2005, pesonal communication; Liste and Liste, 1990). 61

93 Ohio Native Ameicans The sample fom Ohio comes fom a village and cemetey nea Madisonville and was descibed in a book by Hooten and Willoughby (1920). The anatomical analysis suggested a close mophological similaity to the Ioquois but Hooten (1920) concluded that moe eseach on neaby goups would be necessay in ode to assign population affinity. This population most pobably pacticed hoticultue (Willoughby and Hooten, 1920). euvian This is a collection of ten euvian pehistoic mummies fom the coastal Ancon egion in the Lima povince in cental eu. The coastal euvian populations ae believed to have pacticed an intensive agicultual lifestyle (Moseley, 2001). Thee ae two individuals fom Choillos, which is south of Lima and also a coastal aea whee a simila agicultual lifestyle was pacticed (Moseley, 2001). ygmy Fou twentieth-centuy ygmy individuals (pobably Aka) fom Cental Afican Republic ae included in the sample. The tem ygmy as used hee is a deogatoy tem that efes to a shot statued goup of populations fom cental Afica, but no bette name is available to descibe these diffeent tibes of foest living goups. The ygmy have hunte-gathee lifestyles and mainly live in the Afican ainfoest. The moden Aka, compaed to some othe ygmy goups, spend most of thei time in the foest and build semi-pemanent camps whee most of the family esides. Foaging makes up most of the subsistence of this goup, although some meat is acquied though collective net hunting (Bahuchet, 1990; Hewlett, 1996). Russian Eskimo The sample of Sibeian eninsula Eskimo is fom a site in Ekveni. It is believed that the individuals lived a taditional life on the nothen ice caps and along the coast of the Sibeian 62

94 eninsula, elying mainly on fish and sea mammals fo thei subsistence (D. ezhemsky, cuato, 2007 pesonal communication). Russian Mesolithic This Russian Mesolithic sample is cuated is fom a site in Vasilievsky. Vasilievsky is an island in the Baltic Sea and is a distict of Saint etesbug, Russia. An excavation yielded a Mesolithic sample of which 15 specimens wee digitised. The individuals wee unsexed and not soted pe individual. The indigenous people of the aea wee hunte-gathees and may have been seasonally nomadic (pesonal communication, J. Chistov). The close poximity to the sea might have made it possible fo the inhabitants to settle in the egion yea-ound and include a significant amount of fish in thei diet. Sibeia The Sibeian sample is cuated at two museums: the Royal Institute fo Natual Sciences, Bussels and the Museum of Anthopology at the State Univesity of Moscow. The Belgium sample compises two individuals, which wee excavated in Sibestey and wee found in close poximity of each othe. The Russian sample comes fom Evenki in Nothen Tansbaikalia, Sibeia, and ae dated to 1000BC-1000AD. Moden inhabitants of Evenki still pactice a taditional lifestyle, and thee is no eason to believe that this lifestyle was not also chaacteistic of the achaeological peoples. Abe (2005) descibes small semi-pemanent, family goup settlements subsisting on small scale yea ound mammal hunting. They pefeed lage game such as mutton and eindee but hunted othe animals oppotunistically fo the est of the yea (Abe, 2005). South Dakota Native Ameicans The South Dakota Native Ameican sample was found on the Oahe Resevation and is fom afte 1750 AD. They ae most likely Aikaa, although the sample is not assigned to a specific population. The post-contact Aikaa had some sedentay settlements and wee mainly 63

95 equestian hunte-gathees. They wee not pacticing agicultue o hoticultue at these sites (Owsley and Jantz, 1994). Tasmanian Two Tasmanians ae included in the sample. One specimen is cuated at the Royal Institute fo Natual Sciences in Bussels, the othe, until ecently, was cuated at the Natual Histoy Museum in London. The Tasmanians ae the extinct aboiginal population of Tasmania, an island 275 km south of Austalia, and wee a physically distinct population fom the Austalian Aboigines because of the sepaation of Tasmania fom geate Austalia between and 6000 B (Wundely, 1938; Henich, 2004). They had no clothing o contol of fie and the achaeological ecod shows that they stopped eating deep sea fish aound 4000 yeas ago but still ate cayfish and shellfish. They wee mainly hunte-gathees who hunted bids, kangaoo, wallaby and opossum (Wundely, 1938; Henich, 2004). Tiea del Fuego Tiea del Fuego is an achipelago south of the southenmost tip of mainland Agentina. The southenmost point of the Islands is Cape Hon. A ight femu, ulna and adius of a single individual that was descibed as having syphilis in the left side of the body was included along with an isolated femu belonging to a diffeent individual. It is unclea to which of the Feugian goups this mateial belongs. Howeve, the elatively small statue of the individual with the femu, ulna and adius would seem to peclude an Ona affinity while the statue of the isolated femu is consistent with Ona affinity. The Fuegians ae not a homogenous goup but athe thee distinct goups, living on diffeent islands with diffeent languages, diffeent appeaances and diffeent cultues. The fist goup ae the Aliculufs (also Halakwulup o Alacaluf), the second the Yahgans (also Yagan o Yaganes). These two goups ae the most closely elated in appeaance. They ae stocky and shot statued, woe vey little o no clothes, despite the cold weathe conditions, lived in canoes and fed off 64

96 mussels, snails, cabs, and fish. The thid goup is distinctly diffeent. They ae the Ona and ae vey tall and have been descibed as giant Indians. They used no canoes and wee huntegathees (Gusinde, 1939; Bollen, 2000). The Fuegians wee descibed as being mophologically close to Neandethals (Gusinge, 1939; Matin, 1959; Genna, 1930). 65

97 3.2. Methods This poject employs a compaative appoach to assess the pattens of mophological vaiation in hominins in elation to aspects of body size, envionmental and behavioual vaiability. The pimay featues unde consideation hee ae cuvatue and apex of cuvatue of the femu, ulna and adius. Howeve, the collection and analysis of univaiate measuements and othe shape vaiables wee collected and analysed fo two puposes: 1) to aid in the intepetation of cuvatue as pat of the est of the mophology, and 2) to investigate the oveall mophology of each of these bones fo the individuals and goups. The functional significance of long bone cuvatue in humans is not well undestood, and a vaiety of hypotheses have been suggested to explain cuvatue in moden humans and Neandethals (Chapte 2). Each of the hypotheses unde consideation will be consideed independently. Data fo each individual is combined with envionmental, geogaphic and behavioual infomation fo the population opulation data and categoies Time peiod The Euopean sample is divided into fou categoies based on time peiod of the sample, and is egadless of activity patten. These categoies ae: Mesolithic, Neolithic, Medieval and 18 th -19 th centuy Envionmental data A numbe of envionmental vaiables wee collected fo each of the moden human population samples: latitude, tempeatue, ainfall, and altitude. Latitude: Mean latitude of the site at which the emains wee discoveed. Latitude is a good 66

98 poxy fo climate as it shows a stong elationship with both mean annual and effective tempeatue (Rose and Vinicius, 2008). Absolute latitude is used fo investigating the elationship between climate and skeletal vaiables. Tempeatue: Aveage annual tempeatue at the place of oigin is used to ceate a distance (diffeence) matix (data fom Hijmans et al., 2005) in ode to detemine if aveage annual tempeatue and cuvatue ae coelated. Rainfall: A matix simila to that fo annual tempeatue was ceated fo aveage annual ainfall (data fom Hijmans et al., 2005) in ode to detemine if thee is a coelation between aveage annual ainfall and cuvatue. Altitude: Aveage altitude at the place of oigin of the population. A dissimilaity matix fo this vaiable is used to test fo a elationship between diffeences in elevation and cuvatue of the femu because high altitude aeas ae typically hilly o mountainous with complex topogaphy. ossible caveats ae high altitude plains whee little elevation diffeences ae found (data fom Hijmans et al., See also Activity levels and subsistence stategy Although thee is a vaiety of ways to quantify the activity levels involved in subsistence activity o habitual behaviou (see Stock, 2002), osteological museum collections ae often limited to making boad cultual genealizations about habitual behaviou. Theefoe, the confidence that can be had in numeical estimates of numbe of moves a yea, distance used ove the couse of the yea and length of the aveage movement is vey low. Beaing this in mind, the populations wee fist classified into thee boad categoies elated to habitual activity levels. The low activity goup ae those who lived in uban aeas and taded fo thei food in an uban setting: mobility levels and activity levels ae low. The modeate activity goup ae individuals who lived in pemanent settlements and elied on intensive agicultue fo subsistence: mobility levels ae low but activity levels ae geneally high. The high activity goup ae foages (hunte-gathees) but also hoticultualists and pastoalists: mobility levels and activity levels ae high in all of these populations. astoalist communities, such as the Saami, have been 67

99 included into the goup of hunte-gathees in some pevious studies because thei heding lifestyle involves long seasonal migations and thus entails a highe level of mobility than the moe sedentay agicultual populations (eason, 2000b). Within these thee boad activity categoies, the high activity goup was divided into five moe naow subsistence categoies: pedestian, equestian and aquatic foages, hoticultualists and pastoalists. These categoies ae used to test fo diffeences in cuvatue associated with specific habitual subsistence behavious. It is impotant to conside these genealisations, and bea in mind that they may not apply to evey individual in the population Individual data In addition to the categoical data fo populations, individual data and univaiate measuements wee collected fo each specimen. Table 3-4 is a list of data collected fo each individual. Table 3-4 Summay of individual data collected duing this poject. Categoy Desciption lace of oigin lace whee the emains wee found o collected opulation opulation name Age Absolute age fo those known o mean of the estimated age ange Age categoy Young adult: epiphyseal sutues visible Adult: no visible epiphyseal sutues o age-elated pathologies Old adult: mild signs of old age such as osteopoosity, athitis pesent (sevee cases excluded) Sex male o female o unknown Side left o ight 68

100 Univaiate measuements Standad univaiate measuements used in osteometic eseach wee collected using the landmaks (see Appendix 1-Appendix 7). Most of these wee descibed in Matin and Salle (1959). The distances o angles wee calculated fom the 3D coodinates (see below ) using geometic methods o vecto algeba. Some of these measuements may not diectly featue in the analyses because they wee used to calculate indices and atios Bone shape In ode to captue shape of the long bones, geometical mophometics is employed hee (Bookstein, 1991; Adams et al., 2004). The elevant analytical appoaches have been developed by a numbe of authos and ae summaised in O Higgins (2000) and Gunz et al. (2005). Geometic mophometics offe consideable advantages ove linea measuements because esults can be visualised as configuations of landmaks in the oiginal space of the specimens athe than only as seconday plots and diagams. This study also includes the use of semi-landmaks, allowing fo the incopoation of outline and suface infomation. Semilandmaks make it possible to include point and outline infomation in a single analysis and to conside the cuves sepaately o as pat of the whole bone mophology. The data fo each individual ae configuations of homologous landmaks and semilandmaks. Each configuation is patitioned into its size and its shape. Size is epesented in the analysis by centoid size, which is the squae oot of the sum of squaed Euclidean distances fom each landmak to the mean of the landmak coodinates. Shape is epesented by the diffeence in coodinates of coesponding landmaks between specimens. These shape coodinates ae the cuves along the suface of the diaphysis and the epiphyses. Diffeences between the configuations can then be used in multivaiate analysis incopoating othe envionmental and behavioual vaiables o coelated with centoid size to exploe the elationship between shape and size (Bookstein, 1991; Runestad et al., 1993; 69

101 Bookstein, 1996; O'Higgins and Jones, 1998; O'Higgins, 2000; Delson et al., 2001; Lockwood et al., 2002; Adams et al., 2004; Gunz et al., 2004; Macus et al., 2004; Gunz et al., 2005) Equipment and softwae Fo the collection of the landmaks and semi-landmaks a Micoscibe 3DX digitise (Immesion Copoation), a laptop compute, Micosoft Excel and Micoscibe Utility Softwae v.4.0 (MUS v. 4.0) wee used. The digitise includes a fine-tipped o ball-tipped stylus attached to a set of mechanical ams. The tips cannot be used duing the same session as they have diffeent lengths. The digitise measues with an accuacy of 0.23mm (inta-obseve eo is discussed below) and is not sensitive to tempeatue, humidity, atmospheic pessue, o magnetic field. Landmaks ae discete points that wee ecoded individually each time the tip of the stylus is activated. The initial semi-landmaks wee ecoded by placing the tip of the stylus on the stat point and ecoding data continuously (evey 5 mm fo adult) along the length of the desied cuve using the auto-plot function in the Micoscibe Utility Softwae v.4.0. Mathematica 5.1 fo Windows (Wolfam Reseach) is a mathematical softwae pogam used fo pe-teatment of the semi-landmaks. The methods used to do this ae descibed below ( ). Afte teatment of the semi-landmaks the landmak configuations wee impoted into Mophologika 2 (O'Higgins and Jones, 1998) Data acquisition and specimen set-up All bones wee initially placed on an osteometic boad whee the 25%, 50% and 80% levels wee taken and maked with small ound stickes. The bone was then placed in the upight position in a suppot with clamps. Both clamps wee coveed with ubbe mateial to ensue gip and minimal damage to the bone. The distal aticulation was placed on the lowest clamp, ensuing the edges of the aticula suface could still be accessed with the digitize. Fo the ulna and adius an elastic band was used to keep the bone fom moving thoughout the digitising pocess. The poximal end was positioned so that it ested between the finges of the uppe 70

102 clamp. This secued the bone without obstucting any measuements. The uppe clamp was then closed ensuing that the bone was not damaged but well secue (Figue 3-1). Fo the femu some exta points wee maked befoe mounting it into the clamps. These wee the most supeio point on the head of the femu and the most infeio point on the distal condyles. Figue 3-1 Specimen set up fo the femu, adius and ulna using clamps and a test tube stand Landmaks and semi-landmaks Landmak points should be homologous acoss specimens. Geometic homology in mophometics is not the same as biological homology (similaity due to common descent). In its pesent use homology efes to coesponding discete geometic stuctues in diffeent individuals, species o thoughout developmental stages. Landmaks and semi-landmaks ae the epesentations of such stuctues (Gunz et al., 2005). Landmaks have been categoised by Bookstein (Bookstein, 1991). Type I landmaks ae pecise juxtapositions of tissues such as tiple points of sutue intesections. Type II landmaks ae associated with, fo example, the maximum of a cuvatue on local stuctues with a biomechanical implication. Type III landmaks ae extemal points o mathematically constucted points like the endpoints of length, beadth, and popotional levels on a bone (e.g. 80%, 50%, 25%) (Bookstein, 1991). Many stuctues, like the long bone diaphysis, lack pecise landmak positions. oints on cuves, fo example, cannot be said to coespond with the same 71

103 points acoss the sample, except in so fa the cuve itself is the same. Semi-landmaks allow fo sufaces and cuves in between type I, II o III landmaks to be included in the analysis by epesenting pats of biological stuctues that coespond acoss specimens. Thity-seven landmaks and fou cuves compised of semi-landmaks wee collected on the femu; twenty-nine landmaks and two cuves wee collected on the adius; and thity-six landmaks and one cuve was collected on the ulna. A list of landmaks and landmak diagams can be found in the Appendix 1 to Appendix Analytical methods Size adjustment fo the linea measuements Some univaiate measuements wee size adjusted by the calculation of atios o indices (Appendix 1 to Appendix 6) multiplied by 100 to facilitate compaisons. Using indices eliminates the effect of scale on the measuement, although allometic effects ae not estimated ocustes methods Supeimposition methods wee used to egiste landmaks and eliminate vaiation due to oveall size. Geneal ocustes analysis (GA; also efeed to as GLS: Genealised Least Squaes) supeimposes landmaks using least-squaed estimates fo otation and tanslation. Fist, the centoid (squae oot of sum of squaed Euclidean distances fom each landmak to the mean of the landmak coodinates) of each landmak configuation was fitted to the oigin (1st specimen), and configuations wee scaled to a common unit size (Adams et al., 2004; Bookstein, 1991). The landmak configuations wee then otated and tanslated to obtain an optimal o closest fit between all points of the configuation and the oigin (Adams et al., 2004; Bookstein, 1991; Bookstein, 1996; O'Higgins, 2000). This pocess was subsequently epeated fo all othe configuations in ode to compute the mean shape. The squaed oot of the sum of the squae 72

104 coodinate diffeences afte supeimposition is a measue of the diffeences in shape between landmak configuations and is called the pocustes distance (Bookstein, 1996). 73

105 Teatment of semi-landmaks Befoe Geneal ocustes Analysis semi-landmaks must be egisteed so that they ae homologous fo compaison between individuals (following Gunz et al., 2005). Fist, a cubic spline is fitted though the ecoded landmaks, and this cubic spine is esampled evey 1mm. Then a desied numbe of equidistant points ae selected along each of the cuves. To test if the numbe of semi-landmaks impacts epeatability, 10 o 20 semi-landmaks on the femu wee chosen. A small numbe of semi-landmaks (compaed to the infinitely lage numbe of points on the cuve) eases computational demand and is sufficient to descibe femoal cuvatue. An altenative to equidistant points is to slide the desied numbe of landmaks along the tangents to the cuve, but this is unnecessay fo simple cuves (Gunz et al., 2005; Bookstein, 1996). Afte this egistation pocedue the configuations wee expoted into Mophologika 2 (O'Higgins and Jones, 1998) fo futhe analysis, togethe with the othe landmaks ecoded duing data collection incipal component analysis incipal component analysis (CA) employs two o moe obsevations fo each individual, which ae then combined to poduce uncoelated indices that explain diffeent dimensions in the data with fewe vaiables than the oiginal obsevations. These indices (called incipal Components) ae odeed so that the fist explains the lagest amount of vaiation and the second explains the second lagest amount of vaiation, and so on. In geometic mophometics incipal Components Analysis is based on elative waps. Relative waps ae linea combinations of patial waps and thei scoes (Dytham, 1999). The whole ange of waps in geometic mophometics ae deived fom thin-plate spine analysis (Slice, 2005). This is the pojection of the points afte GA on a space that is tangent to Kendall s shape space. The shape space is a genealized cuved space with moe than thee dimensions that can be compaed to the suface of the eath and the set of possible shapes fo any given landmak configuations with the same numbe of landmaks and dimensions 74

106 (Monteio et al., 2000). Hee, the distances between the points between two sets of landmaks (efeed to as the distances between pais of points) appoximate the ocustes distances. The fist landmak configuation is usually the efeence, the goup mean, and the second configuation is the taget. The diffeences between single pais of points ae calculated as the displacements of ight angles out of the plane of the efeence. Those equations ae ecombined to expess the totality of diffeences between the two (Adams et al., 2004; Bookstein, 1991; Mitteoecke et al., 2005; Slice, 2005). The gaphical epesentation of landmak configuations makes it easy to visualise shape diffeences (Lockwood et al., 2002; Slice, 2005). These diffeences ae computed duing CA and epesent the total shape vaiability into un-coelated vaiance-maximising vaiables (also called pincipal components). The pecentage vaiance explained by each of the pincipal components is used to detemine which components to examine (based on a scee plot of eigenvalues). These scoes (C scoes) can then be used as data in multivaiate analyses and combined with othe vaiables (Adams et al., 2004; Bookstein, 1991; Mitteoecke et al., 2005; Slice, 2005). Thee is anothe benefit to the use of the 3D mophometic techniques and this is the possibility to use only patial landmak configuations in the CA. Theefoe, in ode to analyse diffeent anatomical featues sepaately, subsets of landmaks and semi-landmaks can be selected and incipal Component Scoes can be used to epesent a cetain tait, athe than the total bone shape. Visualisations using vecto plots of the shape changes along the incipal Components can then be used to intepet the changes in mophology (Slice, 2005). Subsets of data used in the analysis hee ae descibed in the esults chaptes (Chapte 4 and 5) Inta-obseve eo To test the epeatability of the 3D landmaks themselves, data wee ecoded on thee human skeletons at Univesity College London. Each specimen was measued thee times in one week.the ocustes distances fom GA supeimpositions of the landmak and semi-landmak configuations wee used as a measue of obseve eo (Lockwood et al., 2002). This value inceases with inceasing shape diffeence between two specimens. Also, when epeat measuements fom the same individual ae supeimposed using GA, it is possible to identify 75

107 the landmaks with the geatest eo. Floating landmaks such as the middle of a suface o the individual cuves wee expected to vay most. Eo esults fo the geometic mophometic analysis vaied depending on how many semilandmaks wee chosen. Using 20 semi-landmaks, eo diffeences between the thee epeats (mean diffeence 0.017, n=9 compaisons) wee nealy as geat as vaiation between diffeent specimens (mean diffeence 0.018, n=27 compaisons). Using the 10 semi-landmaks, the mean diffeence between specimens was fo 27 compaisons, and the mean diffeence between epeats was fo nine compaisons. Using 10 semi-landmaks along with fixed landmaks, the diffeence between specimens (mean diffeence 0.045, n=27 compaisons) was geate than vaiation between epeats (mean diffeence 0.017, n=9 compaisons). These positive esults fo a intoduced numbe of semi-landmaks imply that the cuve itself was sufficiently descibed by ten semi-landmaks, and additional landmaks eflected eo such as slight hoizontal movement of the hand when ecoding a cuve down the smooth and featueless anteio suface of the femu. Fo this eason, ten semi-landmaks wee used in all analyses Disciminant function analyses Using SSS v.15 disciminant functions wee calculated using the pincipal component scoes fo goups of individuals. This technique maximizes diffeences between known goups and makes pedictions about individuals fo which the goup is not known (Dytham, 1999). In the analyses, goups wee Neandethals, ealy moden humans and ecent moden humans. Only pincipal components that wee found to explain a substantial amount of vaiation (see Chapte 5) ae consideed fo inclusion (Dytham, 1999; Weave, 2002) Analysis of Vaiance (ANOVA) ANOVA was used to detemine the effect of factos influencing cuvatue. ost-hoc tests wee pefomed to identify diffeences between the samples. The samples wee gouped in categoies (see section 3.2.1). Both a Hochbeg s GT2 (fo vey diffeent sample sizes, Field, 2000, p. 341) and a Games-Howell pocedue (fo small and uneven sample sizes whee homogeneity of vaiance is not assumed fo all samples, Field, 2000 p. 341) wee used in SSS v

108 Mantel test Mantel tests wee used to investigate elationships between mophological and envionmental distances between pais of populations using assage v1 (Rosenbeg, 2001). A distance matix is a way of descibing the diffeence (dissimilaity) between pais of populations. A Mantel test tests the null hypothesis that distances in the fist matix ae independent of distances in the second matix. The statistic used fo the measue of the coelation between the two matices is the eason coelation coefficient. In ode to test the null hypothesis, a andomization pocedue is used which compaes the oiginal value of the coelation coefficient to that found by andomly eallocating the ode of the elements in one of the matices (Manly, 1997). Fo each Mantel test mophological distance matices of C scoes fo cuvatue, apex of cuvatue and the whole bone ae coelated to the distance matices of the envionmental factos: tempeatue, ainfall, altitude. Although a numbe of authos have used Mahalanobis distances (Ackemann, 2002; Havati, 2003a; Havati, 2003b; González-José et al., 2004; Havati and Weave, 2006) this poject uses ocustes distances only as they ae not affected by uneven sample sizes and do not assume simila covaiance stuctues fo all goups (Smith et al., 2007) Othe univaiate analyses Depending on the hypothesis being tested, a vaiety of univaiate statistical analyses wee used, including Student s t-test and eason s coelation analyses. Fo coelations with ontogenic age, a non-paametic Kendall s Tau b was used as not all ages wee epesented and the age of some individuals was detemined fom skeletal makes. Fo the ulna and adius the effect of asymmety was investigated using Student s t-test. The sample was collected using the best peseved side of the skeleton. In samples whee pesevation is good, this esulted in a 50/50 split. In some cases, howeve, one of the sides was unavailable fo eseach. The effect of side was tested using a eason s Chi-Squae test on the ecent moden human sample. Despite the esults being affected by small sample sizes o 77

109 samples with only one side epesented (N<5), in about 50% of cases the test is highly significant (p<0.001) indicating that the sampling of left and ight was not independent. Fo this eason, all analyses on the adius and ulna that wee pefomed on the pooled sample wee conducted also fo the ight side only. If the significance values wee affected, those esults will also be epoted. 78

110 3.3. Ode of analysis fo Chapte 4 The pupose of the esults in Chapte 4 is to test the seies of hypotheses and pedictions set out in Chapte 2. The esults will be pesented fist fo the femu and then fo the lowe am. The ode and potocol of the analyses in both sections is descibed hee. Although multiple tests ae conducted that test fo statistical significance, the Bonfeoni coection was not applied. This is pat of a geneal concen that oveuse of the Bonfeoni method may esult in ovely consevative esults (see Moan, 2003; Nakagawa, 2004). Also, in this wok most of the tests ae pefomed to addess specific pedictions and hypotheses, and the chance of spuious significance is educed. Fo the moe exploatoy pats of the analysis, caution is applied when esults do not fit any a pioi expectation, but at the same time these esults ae highlighted given the geneal lack of detailed pevious wok on these skeletal elements Shape data Initially, ocustes coodinates fo all individuals wee analysed using incipal Components Analysis to patition the total shape vaiability into un-coelated vaiance-maximising vaiables. The pecentage vaiance explained by each of the pincipal components was used to detemine which components to examine, based on whee eigenvalues level off on a scee plot. Gaphical epesentations of landmak configuations ae used to visualise shape diffeences and to match each pincipal component to components of cuvatue o othe aspect of shape vaiation Coelations between shaft shape and univaiate measuements. In ode to identify the covaiates with cuvatue and undestand cuvatue as pat of the est of the anatomy, eason s coelations wee pefomed to look fo covaiates between 1) the univaiate measuements, 2) the univaiate measuements and cuvatue and 3) the univaiate measuements and othe aspects of bone shape. These analyses wee pefomed on the whole 79

111 ecent human sample and on the high-activity categoy because expession of skeletal diffeences is moe ponounced in the latte goup. Some pedictions wee made (see Chapte 2: Hypotheses and pedictions) about the elationship of cuvatue to these univaiate measuements, but most of these coelations wee exploatoy. The following eason s coelations wee pefomed: - Femoal shape with neck-shaft angle, tosion angle, femu length, neck length, shaft shape atios (at subtochanteic, midshaft and subpilastic level), obusticity (distal condyles, midshaft and head). - Radius shape with obusticity (head, midshaft and distal aticulation), adius length, neck-shaft angle, position of the adial tubecle, dosal and lateal subtense, neck length, head shape and midshaft shape. - Ulna shape with maximum length, olecanon size, midshaft shape, adial notch size, tochlea notch oientation, olecanon oientation, coonoid-olecanon atio, length of the ponato cest, position of the bachialis insetion and obusticity Body size Reseach on the use of skeletals element in body size estimation has agued fo the use of lowe limb bone dimensions to pedict body size fo moden humans and fossil hominins (see eview in Ruff, 2000a; Auebach and Ruff, 2004; Ruff et al., 2005). As aticula dimensions ae elatively insensitive to vaiations in the mechanical envionment compaed to diaphyseal beadth (which can ove- o undeestimate body size in populations with diffeent activity levels), the femoal head diamete has often been used (Ruff, 1991; McHeny, 1992; Gine et al., 1995) as has bi-iliac beadth (McHeny, 1992; Ruff et al., 1994). Because it was unknown whethe specimens would have the pelvis peseved and less estimation is involved fo femoal head diamete than fo bi-iliac beadth, femoal head-diamete is used as an indicato fo body size in this study. Absolute femoal head diamete was used hee to investigate the elationship of body size with cuvatue. It is known that obusticity scales with body size (van De Meulen et al., 1993; Ruff, 2000a; Stock, 2002; Stock and feiffe, 2004), and this patten was fist confimed fo this sample. Subsequently, the coelation between cuvatue and body size will be investigated using 80

112 population means fo pincipal components elated to cuvatue. Fo the lowe am, this pat of the analysis is possible only when femoal head diamete of the individual was known Sex In ode to assess sexual dimophism in cuvatue, all individuals of known sex wee compaed using a Student s t-test fo obusticity and cuvatue, as well as fo the othe bone shape Cs and the univaiate femu measuements. If thee is a significant elationship between body size and cuvatue, it is expected that sexual dimophism in cuvatue is at least patly elated to diffeences in body size between males and females. Sex diffeences may also be elated to diffeent bone modelling and emodelling ates in males and females o diffeences in activity levels due to sexual division of labou. Simila pedictions have been made fo obusticity, and the pesent sample will theefoe be analysed fo extenal obusticity (at midshaft, distal and poximal condyles) in ode to detemine whethe the sample follows pattens established peviously fo humans. Because sexual division of labou is moe ponounced as geneal activity levels incease, the tests ae epeated fo the high activity categoy (foages, hoticultualists and pastoalists), modeate activity (intensive agicultualists) and low activity (uban tade) categoy samples sepaately Age In ode to investigate whethe cuvatue deceases with deceasing activity levels though adulthood, a Speaman s ank coelation was pefomed on individuals of known age o estimated age. Speaman s ank coelations wee also used to investigate the effect of inceasing age on othe aspects of shaft and epiphyseal shape and othe univaiate measuements to see if they aid the intepetation of tends obseved in cuvatue. Because age is not known o estimated fo the majoity of the sample, age categoies (See section fo details) wee used to test the pedicted elationship fo the sample as a whole. 81

113 Activity pattens The pupose of these analyses is to investigate if cuvatue is highe in populations with high activity levels. The samples wee divided into thee main categoies based as descibed in section Fo each of these analyses, the high activity categoy was divided into five subsistence categoies to investigate if thee ae diffeences between specific foaging stategies in cuvatue of the lowe am and the femu. Othe shaft shape vaiables and univaiate measuements wee also analysed in ode to test the effect of activity levels on the othe aspects of mophology Climate and latitude Although no diect benefit of having a highe degee of cuvatue in colde climates has been suggested, cuvatue may be a consequence of a cold-adapted body shape. In this analysis, climate and specifically tempeatue is quantified using the latitudinal position of the population (Ruff, 1994b; Rose and Vinicius, 2008). Afte eason s coelations ae pefomed between latitude and cuvatue, the othe shaft shape vaiables and univaiate measuements ae also investigated to detemine the suite of mophological featues which vay in esponse to climatic conditions. The analyses will be epeated fo the high activity categoy because these populations may be moe exposed to tempeatue extemes than ae populations in the modeate and low activity categoies Evolution ove time In ode to test fo changes in cuvatue with inceasing sedentism, the Euopean sample is divided into fou categoies: Mesolithic, Neolithic, Medieval and 18th-19th Centuy. Diffeences in cuvatue ae analysed by means of an ANOVA fo pincipal components epesenting cuvatue and apex of cuvatue. 82

114 Mantel test A Mantel test is used to test fo coelations between envionmental factos (tempeatue, ainfall, and altitude) and cuvatue (degee of cuvatue and apex of cuvatue) and whole bone shape. Five thousand pemutations wee pefomed fo each of the tests Systemic influences To investigate whethe cuvatue is systemic, the sample fo which all thee bones ae epesented was used. A eason s coelation analysis was pefomed on the population means fo the degee of cuvatue elated Cs fo all thee bones. 83

115 CHATER 4. INTRASECIFIC DIFFERENCES IN LONG BONE CURVATURE IN MODERN HUMANS 4.1. Objective Chapte 2 illustated the effects of bone emodelling in esponse to use. Distinctive anatomical featues of the long bones ae modified duing development in ways that optimise stength and adaptability in esponse to diffeent activity levels. Hee, the behavioual and envionmental effects on long bone mophology among moden humans ae exploed with the aim of poviding a context to undestand the fossil populations. In the esults descibed below the abbeviations of the pincipal components (Cs) names ae made up of thee pats. The fist designates the landmak set included in the study (i.e. acuve stands fo anteio cuve). The second designates the sample included (i.e. AMH stands fo all ecent moden humans). The thid is the C numbe (i.e. C2 stands fo the second C) The femu Femu shape pincipal components explained The following analyses ae based on the entie sample of moden humans and the analyses wee caied out using the methodology descibed in Chapte 3. The magnitude and patten of vaiation fo the femoal anteio, posteio, medial and lateal cuves ae visualised using Mophologika. Vaiation in the femoal poximal and distal epiphyses ae analysed in a simila fashion. The cuves ae semi-landmaks on the diaphyseal suface only, wheeas the epiphysis analysis uses fixed landmaks (fo details see Chapte 3: Mateials and Methods). In figues, viewing angles wee chosen to best illustate similaities and diffeences. Fo the cuves, this is in lateal view, unless othewise stated. Aows indicate aeas of change. 84

116 Anteio suface (acuve) The fist fou Cs of the anteio cuve analysis explain 61.9%, 8.49%, 7.06% and 6.33%, espectively, of the vaiation (total 73.9%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. The distibution of populations in Figue 4-1 shows the wide ange of vaiation fo C1 compaed to C2. C1 clealy eflects diffeences in degee of anteoposteio cuvatue o subtense (Figue 4-1 and Figue 4-2a). C2 eflects the position of the apex of cuvatue (Figue 4-1 and Figue 4-2b). C3 is the medial o lateal deviation of the distal end of the cuve in anteio view (Figue 4-2 c). C4 is the degee to which the cuve is mediolateally sinusoidal fom anteio view (Figue 4-2d). Figue 4-1 The fist and second Cs fo the anteio cuve of the femu. All ecent moden human samples. 85

117 a b c d Figue 4-2 Mophological tends fo the anteio cuve of the femu fo all ecent moden humans. a: incipal component1: lateal view. Negative values ae less cuved, positive values ae moe cuved. b: incipal component 2: lateal view. Individuals with negative values have a moe poximal apex of cuvatue, wheeas those with positive values have a moe distal apex of cuvatue. c: incipal component 3: anteio view. Negative values have a distal cuve with medial pojection, wheeas positive values have a lateal pojection of the distal cuve. d: incipal component 4: anteio view. Negative values ae the staightest, wheeas positive values indicate a sinusoidal shape. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C osteio suface (pcuve) The fist fou Cs of the posteio cuve analysis explain 28.7%, 14.5%, 10.5% and 6.38%, espectively, of the vaiation (total 60.08%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. 86

118 The posteio Cs ae vey simila to the anteio cuve. C1 eflects vaiation in the degee of anteio cuvatue (Figue 4-3 and Figue 4-4 a). C2 is the posteio pojection of the poximal end of the cuve (Figue 4-4 b). C3 is elated to the apex of cuvatue (Figue 4-3 and Figue 4-4c). C4 is the diection of the posteio pojection of the distal end of the cuve (Figue 4-4d). opulation distibution fo the degee and apex of cuvatue is shown in Figue 4-3. Figue 4-3 The fist and thid Cs fo the posteio cuve of the femu. All ecent moden human samples. Cs ae explained in Figue

119 a b c d Figue 4-4 Mophological tends fo the posteio cuve of the femu fo all ecent moden humans. a: incipal component 1: lateal view. Negative values have a low and positive values have a high degee of cuvatue. b: incipal component 2: lateal view. ositive values have a less posteioly pojected poximal posteio suface and negative values ae moe posteioly pojected. c: incipal component 3: lateal view. Negative values have a highe apex of cuvatue and positive values have a lowe apex of cuvatue. d: incipal component 4: anteio view. ositive values have a less posteioly pojected distal posteio suface and negative values ae moe posteioly pojected distally. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 88

120 Medial suface (mcuve) The fist thee Cs of the medial cuve analysis explain 49.1%, 17.2%, and 5.52%,,espectively, of the vaiation (total 71.82%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. Distibution of populations is shown in Figue 4-5. attens in the fist two Cs ae simila to those of the anteio cuve. C1 eflects diffeences in degee of anteoposteio cuvatue (Figue 4-5 and Figue 4-6a). C2 is elated to the apex of cuvatue (Figue 4-5 and Figue 4-6b). C3 is the posteio pojection of the distal end of the cuve and the evenness of the cuve (Figue 4-6c). Figue 4-5 The fist and second Cs fo the medial cuve of the femu. All ecent moden human samples. Cs ae explained in Figue 4-6Figue

121 a b c Figue 4-6 Mophological tends fo the medial cuve of the femu fo all ecent moden humans. All in lateal view a: incipal component 1. ositive values have a highe degee of cuvatue compaed to negative values. b: incipal component 2. ositive values have a lowe apex of cuvatue, wheeas negative values have a moe poximal apex of cuvatue. c: incipal component 3. ositive values ae moe flattened off with inceased posteio pojection of the distal cuve, wheeas negative values eflect a shaft suface appoaching an ac of a cicle with a lowe degee of posteio pojection distally. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 90

122 Lateal suface (lcuve) The fist fou Cs of the lateal cuve analysis explain 43.8%, 15.2%, 9.08% and 4.82%,espectively, of the vaiation (total 72.93%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. Distibution of the populations ae shown in Figue 4-7. As in the othe cuves anteio cuvatue is the most impotant facto (C1) (Figue 4-7 and Figue 4-8a). (Figue 4-7 and Figue 4-8a). The othe pincipal components fo the lateal cuve ae the most difficult to intepet. C2 is elated to the staightening of the lateal suface of the femu at the level of the lesse tochante (Figue 4-8b). C3 is elated to the apex of cuvatue and the anteio o posteio oientation of the poximal cuve (Figue 4-7 and Figue 4-8c). C4 is the sinusoidal shape of the lateal suface in anteio view (Figue 4-8d). Figue 4-7 The fist and second Cs fo the lateal cuve of the femu. All ecent moden human samples. Cs ae explained in Figue

123 Figue 4-8 Mophological tends fo the lateal cuve of the femu fo all ecent moden humans. a: incipal component 1: lateal view. ositive values have a highe degee of cuvatue and negative values have lowe degees of cuvatue. b: incipal component 2: lateal view. Negative values have a cuve that appoximates an ac on a cicle, wheeas positive values which have a flattening at the poximal end of the cuve c: incipal component 3: lateal view. Negative values have a lowe apex of cuvatue and moe anteio oientation of the poximal cuve compaed to positive values which have a highe apex of cuvatue and a posteioly oiented poximal cuve. d: incipal component 4: anteio view. ositive values ae the staightest, wheeas negative values have an S-cuve. ositive and negative visualisations coespond to the most exteme positive (ight) and negative (left) C scoes on the scale. 92

124 oximal and distal epiphyses (Epi) The fist five Cs of the epiphyses analysis explain 14%, 9.45%, 7.40%, 4.80% and 4.80%, espectively, of the vaiation (total 39.64%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. C1 eflects diffeences in width of the distal epiphyses and neck-shaft angle (Figue 4-9a). C2 is elated to the oveall width of the femu and its epiphyses (Figue 4-9b). C3 is elated to the width of the distal epiphyses and degee of tosion (Figue 4-9c). C4 is not easily intepeted. The changes along the pincipal component ae vey subtle, and this C will theefoe not be consideed futhe in the subsequent analyses. C5 is elated to the length of the femoal neck (Figue 4-9d). 93

125 a b c d 94

126 Figue 4-9 Mophological tends fo the epiphyses of the femu fo all ecent moden humans. All anteio view. a: incipal component 1. Individuals with negative values have wide distal epiphyses, wide shafts and a smalle neck-shaft angle compaed to those with positive values. b: incipal component 2. Individuals with negative values have naowe epiphyses, heads, neck and poximal shaft compaed to those with positive values. c: incipal component 3. Individuals with negative values have naowe distal epiphyses, and moe tosion compaed to those with positive values. d: incipal component 5. Individuals with negative values have a shote neck compaed to those with positive values. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C Summay Degee of anteio cuvatue is the most impotant C fo all fou cuves (acuveamhc1, pcuveamhc1, mcuveamhc1, lcuveamhc1). This is eflected in the significant coelations between the scoes fo the cuvatue Cs (Table 4-1). Because the cuves ae simila in this espect, only the anteio and posteio cuve will be analysed fo degee of anteio cuvatue. Thee is no coelation between the Cs of the epiphyses and the fou cuvatue Cs. Apex of cuvatue (o the position along the shaft whee the maximum subtense is located) is the majo facto in acuveamhc2, pcuveamhc3, mcuveamhc2, lcuveamhc3. Most of these pincipal components ae significantly coelated, although coelations ae lowe than fo Cs elated to the degee of cuvatue (Table 4-12). AcuveAMHC2 and pcuveamhc3 will be used in futhe analyses to epesent the position of the apex of cuvatue. The othe pincipal components fo each of the fou cuves explain mino vaiation in cuve shape and will be included in the analyses to exploe othe aspects of shaft shape in elation to cuvatue. 95

127 Table 4-1 eason s coelation matix: femoal cuvatue Cs (n= 428). acuamhc1 cuvamhc1 cuvamhc ** <0.001 McuAMHC ** 0.241** <0.001 <0.001 LcuAMHC ** 0.382** <0.001 <0.001 ** Coelation is significant at the 0.01 level (2-tailed). * Coelation is significant at the 0.05 level (2-tailed). McuAMHC ** <0.001 Table 4-2 eason s coelation matix femoal apex of cuvatue Cs (N=428) cuvamhc3 McuAMHC2 LcuAMHC3 ** * acuamhc2 cuvamhc3 McuAMHC ** < ** 0.127** < * Coelation is significant at the 0.01 level (2-tailed). Coelation is significant at the 0.05 level (2-tailed). 96

128 Coelations between Cs and univaiate measuements The pupose of these analyses is to establish covaiates between the shape Cs and univaiate measuements in ode to place cuvatue in the context of the est of the anatomy of the femu. All moden humans The cuvatue Cs vay in thei coelations with the univaiate measuements (Table 4-3). Oveall, cuvatue of the posteio suface is positively coelated with obusticity (head, condyles and midshaft). A ounde midshaft shape (midshaftatio) is coelated with a low degee of anteio cuvatue. A ounde poximal shaft (subtochatio) is coelated with a low degee of posteio cuvatue. The diffeent apex of cuvatue Cs vay in thei coelations (Table 4-4). Neck-shaft angle and tosion angle ae negatively coelated with the position of the apex of cuvatue (acuveamhc2). Robusticity of the condyles is coelated with a lowe apex of cuvatue (EpiAMHC1). Shaft shape at the subpilastic atio is negatively coelated with apex of cuvatue (acuvamh2 and pcuvamhc3). Inceasing epiphyseal obusticity is coelated (headob and condylediamatio) with a moe posteioly pojected poximal posteio suface (pcuvamhc2) (Table 4-5). Tosion angle is positively coelated with a moe flattened off medial suface with inceased distal pojection of the distal cuve (McuveAMHC3). Longe femoa have less flattening off poximally of the lateal suface (this flattening eflects the shote femoal shaft by including the slope towads the lesse tochante) (Table 4-6). 97

129 Table 4-3 eason s coelation matix fo femoal cuvatue Cs and univaiate measuements fo all moden human populations (N=36). acuamhc1 cuvamhc Neck-shaft angle tosionangle subtochatio ** midshaftatio 0.450** subpilatio condylediamatio ** obustindex headob ** necklengthatio ** * Coelation is significant at the 0.05 level (2-tailed). ** Coelation is significant at the 0.01 level (2-tailed). FemLength 98

130 Table 4-4 eason s coelation matix fo apex of cuvatue Cs and univaiate measuements fo all moden human populations (N=36). acuamhc2 cuvamhc Neck-shaft angle ** tosionangle * subtochatio midshaftatio subpilatio * * condylediamatio * obustindex headob necklengthatio * Coelation is significant at the 0.05 level (2-tailed). ** Coelation is significant at the 0.01 level (2-tailed). FemLength 99

131 Table 4-5 eason s coelation matix fo othe shaft shape Cs and univaiate measuements fo all moden human populations (N=36). FemLength Neck-shaft angle tosionangle subtochatio midshaftatio subpilatio condylediamatio obustindex headob necklengthatio * ** acuamhc3 acuamhc4 cuvamhc * * * Coelation is significant at the 0.05 level (2-tailed). Coelation is significant at the 0.01 level (2-tailed). cuvamhc McuAMHC * LcuAMHC * LcuAMHC *

132 Table 4-6 eason s coelation matix fo femoal epiphyses shape Cs and univaiate measuements fo all human populations (N=36). FemLength Neck-shaft angle tosionangle subtochatio midshaftatio subpilatio condylediamatio obustindex headob necklengthatio * ** EpiAMHC1 EpiAMHC2 EpiAMHC * * ** * Coelation is significant at the 0.05 level (2-tailed). Coelation is significant at the 0.01 level (2-tailed). EpiAMHC * opulations with high activity levels only The populations with high activity levels (N=21) ae included in the same analyses and all the moden humans above. Oveall, degee of cuvatue is positively coelated with midshaft and subpilastic shaft shape (fo anteio cuvatue) and obusticity (fo posteio cuvatue) (Table 4-7). The length of the neck is elated to the posteio cuvatue. Anteio cuvatue is elated to a ounde shaft shape at midshaft. The diffeent apex of cuvatue Cs also vay in thei coelations with the univaiate measuements (Table 4-8). As epiphyseal obusticity inceases (headob and condylediamatio), apex of the posteio cuve moves distally. Inceasing epiphyseal obusticity is coelated (headob and condylediamatio) with a moe posteioly pojecting poximal posteio suface (pcuvamhc2) (Table 4-9). The length of the femu is positively coelated with a moe even lateal cuve that does not staighten out at the level of the lesse tochante (lcuveamhc2). Midshaft obusticity is negatively coelated with shaft and epiphyseal width and neck-shaft angle (EpiAMHC1) and with obusticity of the poximal and distal epiphyses (EpiAMHC2) (Table 4-10). 101

133 Table 4-7 eason s coelation matix fo femoal cuvatue and univaiate measuements fo populations with high activity levels (N=21). acuamhc1 cuvamhc Neck-shaft angle tosionangle subtochatio midshaftatio 0.724** < subpilatio 0.540* condylediamatio * obustindex ** headob * necklengthatio * * Coelation is significant at the 0.05 level (2-tailed). ** Coelation is significant at the 0.01 level (2-tailed). FemLength 102

134 Table 4-8 eason s coelation matix fo femoal apex of cuvatue and univaiate measuements fo populations with high activity levels (N=21). acuamhc2 cuvamhc Neck-shaft angle tosionangle subtochatio midshaftatio subpilatio condylediamatio ** obustindex headob * necklengthatio *=Coelation is significant at the 0.05 level (2-tailed). **=Coelation is significant at the 0.01 level (2-tailed). FemLength 103

135 Table 4-9 eason s coelation matix fo othe femoal shaft shape Cs and univaiate measuements fo populations with high activity levels (N=21). FemLength Neck-shaft angle tosionangle subtochatio midshaftatio subpilatio condylediamatio obustindex headob necklengthatio * ** acuamhc3 acuamhc4 cuvamhc * * * Coelation is significant at the 0.05 level (2-tailed). Coelation is significant at the 0.01 level (2-tailed). cuvamhc McuAMHC LcuAMHC * LcuAMHC

136 Table 4-10 eason s coelation matix fo femoal epiphyses shape Cs and univaiate measuements fo populations with high activity levels (N=21). FemLength Neck-shaft angle tosionangle subtochatio midshaftatio subpilatio condylediamatio obustindex headob necklengthatio * ** EpiAMHC1 EpiAMHC2 EpiAMHC * * ** ** Coelation is significant at the 0.05 level (2-tailed). Coelation is significant at the 0.01 level (2-tailed). EpiAMHC

137 Summay Oveall, the anteio and posteio degees of cuvatue ae coelated with diffeent vaiables. Individuals with highe levels of posteio cuvatue have highe levels of obusticity. Individuals with a high degee of anteio cuvatue have a ounde shaft at midshaft. Inceased obusticity of the distal and poximal epiphyses is also coelated with a moe distal apex of cuvatue of the posteio cuve. A moe poximal posteio apex of cuvatue is found with high neck-shaft and tosion angles Factos influencing cuvatue in moden humans The following analyses focus on the elationship between anteio femoal cuvatue and the behavioual, envionmental and biological vaiables that might be expected to influence cuvatue. These coelation analyses test the hypotheses and pedictions pesented in Chapte Body Size The pupose of these analyses is to investigate the coelation between body size and cuvatue. Body size is known to be coelated with diaphyseal vaiables, such as coss-sectional geomety and obusticity (Ruff, 2000a; Stock, 2002; Shackelfod, 2007) and may also have an effect on cuvatue. The elationship between body size and obusticity (subtochanteic, midshaft and subpilastic) and cuvatue is analysed fo the whole sample. Using anteoposteio head diamete as an estimate fo body size (Ruff, 1991; McHeny, 1992; Gine et al., 1995) fo the moden human sample (36 populations) the elationship between body size and obusticity and body size and femoal cuvatue and apex of cuvatue ae investigated. Thee is a significant coelation between body size and the thee diffeent measues of obusticity (Table 4-11). Thee is no coelation between cuvatue and the position of the apex of cuvatue and body size (Table 4-12). 106

138 Table 4-11 eason s coelations fo body size (head diamete) and obusticity of the femu (N=36). HeadAdiamete cuvamhc cuvamhc acuamhc acuamhc * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). Table 4-12 eason s coelations fo body size (head diamete) and obusticity of the femu (N=36). headadiamete condylediamatio * ** obustindex headob Coelation is significant at the 0.05 level (2-tailed). Coelation is significant at the 0.01 level (2-tailed). Summay Thee is no allometic elationship between body size and cuvatue o apex of cuvatue Sex The pupose of these analyses is to investigate the effect of sex on cuvatue and apex of cuvatue as well as othe aspects of bone mophology. Diffeences between males and females can eithe be the consequence of highe body size in males than in females (Student s t-test; t=6.507; <0.001), diffeent bone modelling and emodelling ates in males and females, o due to diffeent loading egimes and activity levels because of sexual division of labou. Cuvatue Although obusticity (midshaft and distal epiphyses) is also elated to A femoal head diamete (body size) (Table 4-13) and males have a lage A femoal head diamete (body size) than females (Student s t-test; t=6.507; <0.001), the analysis above did not find a coelation with 107

139 body size and cuvatue. Fo the whole sample of known sex (N=102 males and 89 females), cuvatue is males is not geate than in females (Table 4-14). Table 4-13 Student s t-test esults fo obusticity in moden human males and females. Sex N male 102 female 89 obustindex male 102 female 89 headob male 102 female 89 * = significant at the 0.05 level. condylediamatio Mean S.D T * * Table 4-14 Student s t-test esults fo femoal cuvatue in moden human males and females. Sex N male 102 female 89 cuvamhc1 male 102 female 89 * = significant at the 0.05 level. acuamhc1 Mean S.D T Fo the samples in this study the pediction that the effect of sex on obusticity and cuvatue is moe evident in goups with high activity levels than in populations with modeate o low activity levels is only patly met. The pediction is met fo two out of thee measues of obusticity fo those with high activity levels (N=41 males and 44 females) and as fo the whole sample, males have highe midshaft and distal epiphysis obusticity (condylediamatio) than females (Table 4-15). Fo the high activity goup the degee of cuvatue is highe in males fo the anteio suface but not fo the posteio (Table 4-18). Fo the modeate activity goup (N=34 males and 28 females), thee is a significant diffeence in midshaft obusticity (Table 4-16) but no diffeence in cuvatue (Table 4-19, Table 4-20). Fo the low activity goup, thee ae no diffeences between males and females in obusticity (Table 4-17) o cuvatue (Table 4-20). In the analysis of the entie human sample the diffeences between males and females with high activity levels ae masked by the similaity between males and females with modeate and low activity levels. 108

140 Table 4-15 Student s t-test esults fo obusticity in moden humans with high activity levels. Sex N condylediamatio male 42 female 44 obustindex male 42 female 44 headob male 42 female 44 * = significant at the 0.05 level. Mean S.D T * * Table 4-16 Student s t-test esults fo obusticity in moden humans with modeate activity levels. Sex N male 34 female 28 obustindex male 34 female 28 headob male 34 female 28 * = significant at the 0.05 level. condylediamatio Mean S.D T <0.001* Table 4-17 Student s t-test esults fo obusticity in moden humans with low activity levels. Sex N male 26 female 18 obustindex male 26 female 18 headob male 26 female 18 * = significant at the 0.05 level. condylediamatio Mean S.D T Table 4-18 Student s t-test esults fo cuvatue in moden humans with high activity levels. Sex N male 42 female 44 cuvamhc1 male 42 female 44 * = significant at the 0.05 level. acuamhc1 Mean S.D T * Table 4-19 Student s t-test esults fo cuvatue in moden humans with modeate activity levels. Sex N male 34 female 28 cuvamhc1 male 34 female 28 * = significant at the 0.05 level. acuamhc1 Mean S.D T

141 Table 4-20 Student s t-test esults fo cuvatue in moden humans with low activity levels. Sex N male 26 female 18 cuvamhc1 male 26 female 18 * = significant at the 0.05 level. acuamhc1 Mean S.D T Apex of cuvatue Fo all individuals (N=102 males and 89 females), females have a lowe apex of cuvatue than males (acuveamhc2; p=0.034) (Table 4-21). This diffeence is not pesent in goups with high activity levels (N=42 males and 44 females) (Table 4-22) o low activity levels (N=26 males and 18 females) (Table 4-24). Only fo goups with modeate activity levels (N=34 males and 28 females) (Figue 4-16 and Table 4-25) is thee a significant diffeence between males and females. Table 4-21 Student s t-test esults fo apex of cuvatue in moden human males and females. Sex N male 102 female 89 cuvamhc3 male 102 female 89 * = significant at the 0.05 level. acuamhc2 Mean S.D T * Table 4-22 Student s t-test esults fo apex of cuvatue in moden humans with high activity levels. Sex N male 42 female 44 cuvamhc3 male 42 female 44 * = significant at the 0.05 level. acuamhc2 Mean S.D T Table 4-23 Student s t-test esults fo apex of cuvatue in moden humans with modeate activity levels. Sex N male 34 female 28 cuvamhc3 male 34 female 28 * = significant at the 0.05 level. acuamhc2 Mean S.D T *

142 Table 4-24 Student s t-test esults fo apex of cuvatue in moden humans with low activity levels. Sex N male 26 female 18 cuvamhc3 male 26 female 18 * = significant at the 0.05 level. acuamhc2 Mean S.D T Othe shaft shapes Males have significantly staighte poximal posteio diaphyses wheeas those of females slope posteioly (pcuveamhc2, Student s t-test, p=0.031) (Table 4-25). Table 4-25 Student s t-test esults fo othe aspects of shaft shape in moden human males and females. Sex N male 102 female 89 acuamhc4 male 102 female 89 cuvamhc2 male 102 female 89 cuvamhc4 male 102 female 89 McuAMHC3 male 102 female 89 LcuAMHC2 male 102 female 89 LcuAMHC4 male 102 female 89 * = significant at the 0.05 level acuamhc3 Mean S.D T * Epiphysis mophology Males and females ae simila in thei epiphyseal mophology. None of the Cs show distinct diffeences between males and females fo the whole sample (Table 4-26). Thee is only one significant sex diffeence fo the subsample with low activity levels fo EpiAMHC3 (Table 4-27; Table 4-28; Table 4-29). This suggests that males have wide distal condyles and moe tosion than females with low activity levels. 111

143 Table 4-26 Student s t-test esults fo epiphysis shape in moden human males and females. Sex N Mean male female EpiAMHC2 male female EpiAMHC3 male female EpiAMHC5 male female * = significant at the 0.05 level. EpiAMHC1 S.D t Table 4-27 Student s t-test esults fo epiphysis shape in moden humans with high activity levels. Sex N Mean male female EpiAMHC2 male female EpiAMHC3 male female EpiAMHC5 male female * = significant at the 0.05 level. EpiAMHC1 S.D T Table 4-28 Student s t-test esults fo epiphysis shape in moden humans with modeate activity levels. Sex N Mean male female EpiAMHC2 male female EpiAMHC3 male female EpiAMHC5 male female * = significant at the 0.05 level. EpiAMHC1 S.D t Table 4-29 Student s t-test esults fo epiphysis shape in moden humans with low activity levels. Sex N Mean male female EpiAMHC2 male female EpiAMHC3 male female EpiAMHC5 male female * =significant at 0.05 level. EpiAMHC1 S.D t *

144 Univaiate measuements Males have significantly longe femoa in the combined moden human sample (Table 4-30) and in each of the thee activity subsamples (Table 4-31; Table 4-32; Table 4-33). Males with high activity levels also have a ounde distal shaft (subpilatio) than females. Table 4-30 Student s t-test esults fo univaiate measuements in moden human males and females. Sex N male 102 female 89 Neck-shaft angle male 102 female 89 Tosion angle male 102 female 89 subtochatio male 102 female 89 midshaftatio male 102 female 89 subpilatio male 102 female 89 * = significant at the 0.05 level. Femu length Mean S.D t <0.001* * Table 4-31 Student s t-test esults fo univaiate measuements in moden humans with high activity levels. Sex N male 42 female 44 Neck-shaft angle male 42 female 44 Tosion angle male 42 female 44 subtochatio male 42 female 44 midshaftatio male 42 female 44 subpilatio male 42 female 44 * = significant at the 0.05 level. Femu length Mean S.D t <0.001* * 113

145 Table 4-32 Student s t-test esults fo univaiate measuements in moden humans with modeate activity levels. Sex N male 34 female 28 Neck-shaft angle male 34 female 28 Tosion angle male 34 female 28 subtochatio male 34 female 28 midshaftatio male 34 female 28 subpilatio male 34 female 28 * = significant at the 0.05 level. Femu length Mean S.D t * Table 4-33 Student s t-test esults fo univaiate measuements in moden humans with low activity levels. Sex N male 26 female 18 Neck-shaft angle male 26 female 18 Tosion angle male 26 female 18 subtochatio male 26 female 18 midshaftatio male 26 female 18 subpilatio male 26 female 18 * = significant at the 0.05 level. Femu length Mean S.D t * Summay Fo the whole sample males have longe and moe obust femoa than females. Males also have elatively wide knees and staighte poximal anteio shafts. Males do not have highe levels of cuvatue when the whole ecent moden human sample is consideed. Theefoe, cuvatue is not due to diffeences in bone modelling and emodelling between males and females. The pediction that males would have a highe degee of cuvatue and highe obusticity due to having highe activity levels is suppoted fo goups with high activity levels in which division of labou is moe ponounced. 114

146 Age The pupose of these analyses is to investigate the changes in femoal cuvatue thoughout adulthood. If cuvatue is a plastic featue that esponds to habitual loading, it is pedicted that as activity levels decease with inceasing age (fo this skeletally adult sample up to 87 yeas old), so will degee of cuvatue. Of the whole sample, only 88 individuals had known ages o age ange estimates (epesented populations: Afican-Ameican, Aleut, Andamanese, Austalians, English Medieval and 18th 19th Centuy, Ohio Native, Natufian, Danish Medieval and Czech Medieval. These populations epesent all thee activity goups). Thee is no elationship between age afte adulthood and cuvatue no is thee a elationship with apex of cuvatue o the othe Cs (Table 4-34). When the univaiate measuements ae compaed to age thee ae thee significant tends visible: age is negatively coelated with tosion and neck-shaft angle and positively coelated with obusticity of the distal condyles (condylediamatio). Olde individuals have wide knees elative to shaft length, lowe femoal tosion and lowe neck-shaft angles (Table 4-35). Table 4-34 Kendall s Tau b coelations fo Cs and age (N=88). Cuvatue acuamhc1 cuvamhc1 Othe shaft shape Cs acuamhc3 acuamhc4 cuvamhc2 cuvamhc4 Apex of cuvatue acuamhc2 cuvamhc3 McuAMHC McuAMHC4 LcuAMHC2 LcuAMHC *=Coelation is significant at the α=

147 Table 4-35 Kendall s Tau b coelations fo univaiate measuements and age (N=88). Femu length Neck-shaft angle <0.001* Tosion angle * subtochatio midshaftatio *=Coelation is significant at the α=0.05 subpilatio condylediamatio necklengthatio obustindex headob * When age categoies (see Chapte 3 fo moe infomation) wee used instead of absolute age of the individual, the ANOVA showed no significant diffeence between the goups (Table 4-36). Table 4-36 ANOVA esults fo adult age categoies on cuvatue Cs (N=4) F acuamhc cuvamhc *=significant at α=0.05 Sig Summay The pediction is not met. Thee is no tend towads lowe degees of cuvatue with inceasing age. Neck-shaft angle and tosion angle decease with inceasing age and the elative size of the distal condyles inceases Activity levels The pupose of the following analyses is to detemine if thee ae diffeences in degee and apex of cuvatue between samples with diffeent activity levels, using the activity goups and subsistence categoies descibed in Chapte 3 and summaised in Appendix

148 Figue 4-10 Distibution of the activity level categoies in the space of C1 (degee of cuvatue) and C2 (apex of cuvatue) of the anteio cuve fo all moden humans. Cicles: high activity; squaes: modeate activity; cosses: low activity. 117

149 Figue 4-11 Distibution of the activity level categoies in the space of C1 (degee of cuvatue) and C2 (apex of cuvatue) of the posteio cuve fo all moden humans. Cicles: high activity; squaes: modeate activity; cosses: low activity. Cuvatue The activity goups ae significantly diffeent in anteio but not in posteio cuvatue (Table 4-37). Fo the two cuvatue elated Cs, those with high activity levels ae the most cuved and those with low activity levels ae the least cuved (Figue 4-12) (Appendix 12). Howeve, the pincipal souce of vaiation is the diffeence between low activity populations and all othes. 118

150 Table 4-37 ANOVA esults fo activity levels and femoal cuvatue Cs. d.f.=2 F acuamhc cuvamhc *=significant at α=0.05 Sig * Figue 4-12 Anteio femoal cuvatue fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Fo the subsistence categoies, thee ae significant diffeences in the degee of posteio cuvatue (cuamhc1) (Table 4-38). The pastoalists have a highe degee of posteio cuvatue than all othe categoies (Figue 4-13) (Appendix 13). Table 4-38 ANOVA esults fo high activity subsistence categoies and femoal cuvatue Cs. d.f.=5 acuamhc1 cuvamhc1 *=significant at α=0.05 F Sig * 119

151 Figue 4-13 osteio femoal cuvatue fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). Apex of cuvatue The activity goups ae significantly diffeent fo apex of cuvatue in both Cs (Table 4-39). ost-hoc pocedues show that high and low activity categoies ae diffeent fom each othe. Low activity goups have the most poximal apex of cuvatue, high acitivity goups the most distal and modeate activity goups ae intemediate (Figue 4-14 and Figue 4-15) (Appendix 14). Table 4-39 ANOVA esults fo activity levels and the apex of femoal cuvatue Cs. d.f.=2 acuamhc2 cuvamhc3 *=significant at α=0.05 F Sig. <0.001* <0.001* 120

152 Figue 4-14 Anteio femoal apex of cuvatue fo moden humans, by activity level. Scale is evesed so that highe values indicate a moe poximal apex of cuvatue. Mean and 95% confidence inteval (whiskes). Figue 4-15 osteio femoal apex of cuvatue fo moden humans, by activity level. Scale is evesed so that highe values indicate a moe poximal apex of cuvatue. Mean and 95% confidence inteval (whiskes). 121

153 The anteio apex of cuvatue is significantly diffeent fo the high activity subsistence goups (Table 4-40). ost-hoc pocedues show that the equestian foages have the most distal apex of cuvatue. The aquatic foages have the most poximal apex of cuvatue and ae significantly diffeent fom the equestian foages and (Appendix 15) (Figue 4-16). Table 4-40 ANOVA esults fo subsistence categoies and the apex of femoal cuvatue Cs. d.f.=5 F acuamhc2 cuvamhc3 *=significant at α=0.05 Sig * Figue 4-16 Anteio femoal apex of cuvatue fo moden humans, by subsistence stategy. Scale is evesed so that highe values indicate a moe poximal apex of cuvatue. Mean and 95% confidence inteval (whiskes). 122

154 Othe elements of shaft shape Fou of the othe shaft shape Cs (pcuveamhc4, mcuveamhc3, lcuveamhc2 and LcuveAMHC4) ae affected by activity level (Table 4-41) (Figue 4-2, Figue 4-4, Figue 4-6, Figue 4-8). ost-hoc pocedues show that the distal end of the diaphysis in the modeate activity goups is staighte, wheeas in high activity goups it is moe posteioly pojected distally (Appendix 15). This could be an indication of moe posteio expansion of the distal condyles (pcuveamhc4) (Figue 4-17). The modeate activity goups also have a moe even cuve that appoximates and ac of a cicle with less posteio pojection of the distal medial suface compaed to the high activity level goups who have a moe flattened off medial cuve with inceased posteio pojection distally (mcuveamhc3) (Figue 4-18). The high and modeate activity goups have a staightening of the femu at the level of the lesse tochante, wheeas those with low activity levels have a lateal suface that appoximates the suface of a cicle (lcuveamhc2) (Figue 4-19). The low activity populations ae also significantly diffeent (<0.001) in having a lateal suface that, in anteio view, is sinusoidally shaped, wheeas high and modeate activity goups have a moe even lateal suface (lcuveamhc4) (Figue 4-20). 123

155 Table 4-41 ANOVA esults fo activity levels and the othe femoal shaft shape Cs. d.f.=2 acuamhc3 acuamhc4 cuvamhc2 cuvamhc4 McuAMHC3 LcuAMHC2 LcuAMHC4 *=significant at α=0.05 F Sig <0.001* <0.001* 0.001* <0.001* Figue 4-17 cuvamhc4 fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Negative values have a posteio expansion of the distal epiphyses eflecting moe posteioly pojecting condyles. 124

156 Figue 4-18 McuvAMHC3 fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). High values have a moe flattened off medial cuve with inceased posteio pojection distally. Figue 4-19 LcuvAMHC2 fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Low values have a lateal suface that appoximates an ac of a cicle. 125

157 Figue 4-20 LcuvAMHC4 fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Lowe values have a lateal suface that in anteio view is sinusoidally shaped. 126

158 The same analysis fo the othe shaft shape Cs was epeated fo the high activity subsistence goups (Table 4-42). Thee ae significant diffeences between the goups fo pcuveamhc2 and post-hoc compaisons indicate that equestian foages have a significantly poximally staighte posteio diaphyseal suface compaed to aquatic foages and pastoalists (Figue 4-21 and Appendix 17). Table 4-42 ANOVA esults fo subsistence categoies and the othe femoal shaft shape Cs. d.f.=5 acuamhc3 acuamhc4 cuvamhc2 cuvamhc4 McuAMHC3 LcuAMHC2 LcuAMHC4 *=significant at α=0.05 F Sig * Figue 4-21 cuvamhc2 fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). High values have a poximally staighte posteio diaphyseal suface. 127

159 Univaiate measuements The activity goups ae significantly diffeent in femu length, neck-shaft angle, tosion, shaft shape at the sub-tochanteic and sub-pilastic level, neck length atio and obusticity of the shaft and head (Table 4-43, Figue Figue 4-29). High activity goups have significantly moe obust and shote femoa with a highe neck-shaft angle and a shote neck than low activity goups (Appendix 18) (Figue Figue 4-29). Modeate activity goups also have a longe femu than high activity goups (Figue 4-22). The low activity goups have ounde shafts at the sub-tochanteic and sub-pilastic level and ae anteoposteioly wide at the midshaft level compaed to high and modeate activity goups (Figue Figue 4-26). Table 4-43 ANOVA esults fo activity level and the univaiate measuements of the femu. d.f.=2 Femu length Neck-shaft angle Tosion angle subtochatio midshaftatio Subpilatio condylediamatio necklengthatio obustindex Headob *=significant at α=0.05 F Sig. <0.001* <0.001* <0.001* 0.028* <0.001* <0.001* 0.002* 0.002* 128

160 Figue 4-22 Femu length fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Figue 4-23 Neck-shaft angle fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). 129

161 Figue 4-24 Subtochanteic shape atio fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Figue 4-25 Midshaft shape atio fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). 130

162 Figue 4-26 Subpilastic shape atio fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Figue 4-27 Neck-length atio fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). 131

163 Figue 4-28 Robusticity index fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Figue 4-29 Femoal head obusticity fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). 132

164 The subsistence categoies with high activity levels ae also diffeent fo some of the univaiate measuements (Table 4-44). Most univaiate measuements have significant between goup diffeences, with the exception of sub-tochanteic and sub-pilastic shaft shape. The post-hoc pocedues (Appendix 19) show that the aquatic foages have shote femoa than all othe goups (not statistically significant fo hoticultualists) (Figue 4-30). Equestian foages and pastoalists have lowe neck-shaft angles (Figue 4-31). Equestian foages have the lowest amount of femoal tosion and the smallest femoal head size (Figue 4-32, Figue 4-33). Thee is a tend fom anteoposteioly wide to ound shafts though the diffeent subsistence stategies, but not all goups ae significantly diffeent fom each othe. This tend may eflect changes in the anatomy with the adoption of subsistence stategies with lowe activity intensity (Figue 4-33). astoalists have the highest obusticity indices at midshaft (Figue 4-34). Table 4-44 ANOVA esults fo subsistence categoies and the femoal univaiate measuements. d.f.=5 Femu length Neck-shaft angle Tosion angle subtochatio midshaftatio Subpilatio condylediamatio necklengthatio Robustindex Headob *=significant at α=0.05 F Sig. <0.001* <0.001* 0.001* * <0.001* <0.001* <0.001* 0.028* 133

165 Figue 4-30 Femu length fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). Figue 4-31 Neck-shaft angle fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). 134

166 Figue 4-32 Tosion angle fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). Figue 4-33 Midshaft shape fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). 135

167 Figue 4-34 Robusticity index fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). Figue 4-35 Femoal head obusticity fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). 136

168 Epiphysis shape The activity level goups ae significantly diffeent fo 2 out of 5 epiphysis shape Cs (EpiAMHC2 and EpiAMHC5) (Table 4-45; Figue 4-36 and Figue 4-37). Goups with high activity levels have wide epiphyses compaed to goups with modeate o low activity levels, and both high and modeate activity goups have a shote neck than goups with low activity levels (Appendix 20). Fo the subsistence goups, 3 out of 4 Cs wee significantly diffeent: EpiAMHC1, EpiAMHC3 and EpiAMHC5 (Table 4-46). astoalists have wide epiphyses than hoticultualists, pedestian and aquatic foages. The equestian foages ae intemediate (EpiAMHC2) (Figue 4-38). edestian foages have less tosion and have wide distal epiphyses than equestian foages and pastoalists but ae not diffeent fom the aquatic foages and hoticultualists (EpiAMHC3) (Figue 4-39). Aquatic foages have a longe neck than pedestian and equestian foages. The othe categoies ae not diffeent fom each othe (EpiAMHC5) (Figue 4-40) (Appendix 21). Table 4-45 ANOVA esults fo activity level categoies and the femoal epiphyses shape Cs. d.f.=2 EpiAMHC1 EpiAMHC2 EpiAMHC3 EpiAMHC5 *=significant at α=0.05 F Sig * * 137

169 Figue 4-36 EpiAMHC2 (epiphysis width) fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Figue 4-37 EpiAMHC5 (neck length) fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). 138

170 Table 4-46 ANOVA esults fo subsistence categoies and the femoal epiphyses shape Cs. d.f.=5 EpiAMHC1 EpiAMHC2 EpiAMHC3 EpiAMHC5 *=significant at α=0.05 F Sig. <0.001* <0.001* <0.001* Figue 4-38 EpiAMHC1 (epiphysis width) fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). 139

171 Figue 4-39 EpiAMHC3 (tosion and distal epiphysis width) fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). Figue 4-40 EpiAMHC5 (neck length) fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). 140

172 Summay As pedicted, the high activity goup has a highe degee of cuvatue and a lowe apex of cuvatue than the low activity goup. The modeate activity goup is intemediate and significantly moe cuved that the low activity goup. Within the high activity goups the pastoalists wee the most cuved. This may eflect thei highe levels of teestial mobility compaed to the othe high activity categoies. Aquatic foages have the highest apex of cuvatue. This could be a eflection of thei pefeence fo the use of watecaft fo subsistenceelated activity and eflect the esulting educed amount of teestial mobility. Inceased cuvatue fo the high activity goups coincides with inceased obusticity, a moe mediolateally wide shaft and a shote femoal neck Evolution ove time in Euope The pupose of the following analyses is to detemine if, with time, pattens of cuvatue have been affected by the adoption of inceasingly sedentay lifestyles in Euope (Appendix 8). Cuvatue The pediction is that Euopean populations fom the Mesolithic though to the 18th-19th centuy would show deceasing degees of cuvatue (Figue 4-41 and Figue 4-42). Thee is no significant diffeence between time peiods in Euope (Table 4-47). The degee of anteio cuvatue does decease (Figue 4-43), but the posteio cuve shows a diffeent patten (Figue 4-44 and Figue 4-45). 141

173 Figue 4-41 Distibution of time peiods in the space of C1 (degee of cuvatue) and C2 (apex of cuvatue) of the anteio cuve fo all moden humans. Cicles: 18th-19th C, tiangles: Medieval, squaes: Mesolithic, cosses: Neolithic. 142

174 Figue 4-42 Distibution of the time peiods in the space of C1 (degee of cuvatue) and C3 (apex of cuvatue) of the posteio cuve fo all moden humans. Cicles: 18th-19th C; tiangle: Medieval; squaes: Mesolithic; cosses: Neolithic. Table 4-47 ANOVA esults fo time peiod and the femoal cuvatue Cs. d.f.=3 acuamhc1 cuvamhc1 *=significant at α=0.05 F Sig

175 Figue 4-43 Anteio femoal cuvatue fo moden Euopeans, by time peiod. Mean and 95% confidence inteval (whiskes). Figue 4-44 osteio femoal cuvatue fo moden Euopeans, by time peiod. Mean and 95% confidence inteval (whiskes). 144

176 Apex of cuvatue Time peiod has a significant effect on the anteio apex of cuvatue (Table 4-48). ost-hoc compaisons show that 18th-19th centuy samples have a highe apex of cuvatue compaed to Medieval samples but ae not significantly diffeent fom the othe populations (Appendix 22) (Figue 4-45). Table 4-48 ANOVA esults fo time peiod and the femoal cuvatue Cs d.f.=3 acuamhc2 cuvamhc3 *=significant at α=0.05 F Sig * Figue 4-45 Anteio apex of femoal cuvatue fo moden Euopeans, by time peiod. Mean and 95% confidence inteval (whiskes). Summay The pediction is not suppoted because no significant diffeences between the time peiods wee found. The plots, howeve, show that with inceasing sedentism thee is a deceasing tend in degee of anteio cuvatue. The posteio cuve follows an opposite patten, howeve. This may be because time peiod does not accuately eflect a decease in activity levels and loading. 145

177 Climate and latitude As discussed in Chapte 3 latitude is used hee as a geneal poxy fo climate (Appendix 8). Thee is no coelation between latitude and cuvatue, o apex of cuvatue (Table 4-51). Individuals in highe latitudes have wide poximal and distal epiphyses (EpiAMHC1) (Table 4-49). The elationship between the univaiate measuements and latitude ae also investigated and follow peviously established pattens (Table 4-50). Individuals living in highe latitudes have highe levels of obusticity and a elatively longe femoal neck. Thee was a positive coelation between femu length and latitude. This elationship was supising but when the data wee investigated, the coelation appeaed skewed by small-bodied populations living in low latitudes (Figue 4-46) and by the lack of tall equatoial goups in the sample. When the Andamanese, ygmy and euvian ae excluded the coelation is negative, but not significant (=-0.353; =0.051; N=31). 146

178 Table 4-49 eason s coelations fo cuvatue, apex of cuvatue, diaphyseal shape and epiphyses shape Cs and latitude (climate) on the femu (N=35). Absolute latitude (N=35) Cuvatue cuvamhc1 acuamhc Othe shaft shape acuamhc3 acuamhc4 cuvamhc2 cuvamhc4 Apex of cuvatue acuamhc2 cuvamhc McuAMHC3 LcuAMHC2 LcuAMHC4 * ** Coelation is significant at the 0.05 level (2-tailed). Coelation is significant at the 0.01 level (2-tailed) Epiyphyses EpiAMHC1 EpiAMHC2 EpiAMHC3 EpiAMHC4 EpiAMHC **

179 Table 4-50 eason s coelations fo femoal univaiate measuements and latitude (climate) on the femu (N=35). Absolute latitude Univaiate measuements FemLength 0.354* subpilatio Neck-shaft angle * condylediamatio tosionangle necklengthatio subtochatio obustindex midshaftatio headob * Coelation is significant at the 0.05 level (2-tailed). ** Coelation is significant at the 0.01 level (2-tailed) * ** ** ** Figue 4-46 Femu length and absolute latitude fo the ecent moden human sample including the small bodied equatoial samples: ygmy, euvian and Andamanese samples. Femu shape Cs fo goups with high activity levels only Because the high activity goups have moe anteioly cuved femoa than the modeate and low activity levels and the high activity goups ae possibly moe exposed to climatic conditions without pemanent housing and insulation, the coelations wee epeated fo the high activity 148

180 goups. The coelation between epiphysis width and neck-shaft angle stands (EpiAMHC1). Thee is still no coelation between degee and apex of cuvatue and latitude. A positive coelation exists with LcuAMHC2, indicating that individuals living in highe latitudes have femoa that ae staighte at the level of the lesse tochante (Table 4-51). Table 4-51 eason s coelations fo cuvatue, apex of cuvatue, diaphyseal shape and epiphyses shape Cs and latitude (climate) on the femu in high activity goups (N=17). Absolute latitude Cuvatue acuamhc1 cuvamhc1 Apex of cuvatue acuamhc cuvamhc4 McuAMHC3 LcuAMHC2 LcuAMHC4 * Coelation is significant at the 0.05 level (2-tailed). ** Coelation is significant at the 0.01 level (2-tailed). cuvamhc Othe shaft shape acuamhc3 acuamhc4 cuvamhc * Epicondyles EpiAMHC1 EpiAMHC2 EpiAMHC3 EpiAMHC4 EpiAMHC *

181 Mantel test The Mantel tests take a diffeent appoach fom the latitude analysis, in this case compaing envionmental diffeences to shape diffeences. Results ae summaised in Table Thee is a significant coelation between anteio femoal cuvatue (acuvec1 distances) and altitude diffeences. No coelation exists between cuvatue (acuvec1 distances), apex of cuvatue (acuvec2 distances) o the whole femu shape (includes all Cs used in the analyses above) and aveage ainfall and aveage tempeatue diffeences. Table 4-52 Results of the Mantel tests pefomed fo envionmental distance matices - femu Ant. cuvatue apex of cuvatue all femu Cs altitude * ainfall tempeatue = eason coelation coefficient. All pobabilities based on 5000 pemutations Summay Cuvatue Thee is no coelation between body size and femoal cuvatue so the pediction that cuvatue would be elated to body size was not met. Shaft shape and measues of extenal obusticity ae covaiates of anteoposteio femoal cuvatue. Individuals with a highe degee of cuvatue have highe obusticity levels and ae moe anteoposteioly wide. This suppots the pediction that degee of cuvatue and obusticity ae elated. Anteio cuvatue does not elate to climate but is a good indicato of activity levels. Goups with high activity levels ae the most cuved and, among them, especially those with high levels of teestial mobility (pastoalists). Goups with low activity levels ae the least cuved. Aquatic foages ae less cuved than the othe high activity goups. This is in suppot of the pediction made in Chapte 2. Altitude diffeences ae coelated with anteio cuvatue diffeences which 150

182 may suppot the impotance of the effect of teestial mobility ove subsistence elated activity. Thee is no diffeence between all males and females, and thee is no coelation between body size and degee of cuvatue. Theefoe, the obseved diffeences in degee cuvatue between males and females in high activity goups eflects sexual division of labou, athe than sex diffeences in body size o bone modelling and emodelling ates. Apex of cuvatue Apex of cuvatue is also not elated to body size o climate. Thee is some indication that highe levels of extenal obusticity and a moe anteoposteioly wide shaft ae elated to a moe distal apex of cuvatue. Apex of cuvatue is a good indicato of activity levels. Goups with high activity levels have a moe distal apex. Goup with low activity levels have the most poximal apex of cuvatue. Among the high activity level subsistence stategies, aquatic foages have the most poximal apex of cuvatue and the equestian foages the most distal. Rest of the mophology No pedictions wee made about othe aspects of shaft shape and the univaiate measuements; howeve, thee wee some inteesting esults. The low activity goup had a lateal suface that appoximated an ac of a cicle moe and a lateal suface that, fom an anteio view, was moe sinusoidal than modeate and high activity goups who show a staightening of the poximal lateal suface. The equestian foages stood apat fom the othe subsistence categoies in having a poximally staighte posteio diaphyseal suface compaed to aquatic foages and pastoalists. Thee is no elationship between climate and femoal shaft shape but individuals fom colde aeas do have geate epiphyseal obusticity. The high activity goups had moe obust femoa (at midshaft and epiphyseal) with a highe neck-shaft angle and elatively longe femoal neck length. Low activity goups had moe anteoposteioly wide femoal shafts compaed to high and modeate activity goups who wee not diffeent fom each othe. Among the high activity goups the equestian foages and pastoalists had the lowest neck-shaft angles and in the pastoalists this was combined with high levels of midshaft obusticity. 151

183 4.3. The lowe am As fo the femu, the analyses on the lowe am ae based on the entie sample of moden humans and the Cs fo the adius and ulna will be pesented fist with thei espective visualisations. Subsequently, the esults ae pesented in the same ode as the femu Radius pincipal components explained Medial suface (mcuve) The fist thee Cs of the medial cuve explain 40.4%, 19.7% and 8.75%,,espectively, of the vaiation (total 68.9%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. Figue 4-47 shows the distibutions fo the populations fo C1 and C2. C1 eflects the diffeences in lateal cuvatue of the inteosseous cest (Figue 4-47 and Figue 4-48a). C2 is elated to the medial expansion of the poximal inteosseous cest and the mediolateal diection of the distal end of the medial suface (Figue 4-47 and Figue 4-48b). C3 is the sinusoidal shape of the medial suface in the anteoposteio plane (Figue 4-48c). 152

184 Figue 4-47 The fist and second Cs fo the medial cuve of the adius. All ecent moden human samples. Cs ae explained in Figue

185 a b c Figue 4-48 Mophological tends fo the medial cuve of the adius fo all ecent moden humans. a: incipal component 1: anteio view. Negative values ae moe cuved. b: incipal component 2: anteio view. Negative values show an inceased medial extension of the poximal inteosseous cest and a medial diection of the distal cuve (moe medially expanded ulna notch), wheeas positive values show no medial expansion of the inteosseous cest and an ulna notch that is not medially pojected. c: incipal component 3: medial view. Negative values have a moe sinusoidal shape than positive values which ae staighte. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 154

186 Lateal suface (lcuve) The fist thee Cs of the lateal cuve explain 40.4%, 19.7% and 8.75%, espectively, of the vaiation (total 68.9%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. As fo the medial cuve, C1 of the lateal suface eflects diffeences in lateal cuvatue (Figue 4-49 and Figue 4-50a). The lateal cuve is not affected by the development of the inteosseous cest and can give a bette indication of an apex of cuvatue fo the adius. C2 is influenced by the position of the apex of cuvatue and the diection of the distal end of the lateal suface (Figue 4-49 and Figue 4-50b). C3 elates to the sinusoidal shape of the lateal cuve in the anteoposteio plane (Figue 4-50c). Figue 4-49 The fist and second Cs fo the lateal cuve of the adius. All ecent moden human samples. Cs ae explained in Figue

187 a b c Figue 4-50 Mophological tends fo the lateal cuve of the adius fo all ecent moden humans. a: incipal component 1: antio view. Negative values have a highe degee of cuvatue wheeas positive values have a lowe degee of cuvatue. b: incipal component 2: anteio view: ositive values have a moe poximal apex of cuvatue and a moe lateally pojecting styloid pocess, wheeas negative values have thei apex of cuvatue at midshaft and lack the lateal pojection of the styloid pocess.c: incipal component 3: lateal view. Negative values ae moe sinusoidal compaed to positive values. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 156

188 Epiphyses (Epi) The fist 2 Cs of the epiphysis analysis explain 34.8% and 8.89%,,espectively, of the vaiation (total 43.7%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. C1 eflects the diection of the head and the distal aticula suface in elation to the shaft (Figue 4-51a). C2 elates to the length of the adial neck between the adial tubeosity and 80% level of the shaft and the oientation of the tip of the styloid pocess (Figue 4-51b). 157

189 a b Figue 4-51 Mophological tends fo the epiphyses of the adius fo all ecent moden humans. All medial view. a: incipal component 1. Individuals with negative values have a moe anteioly oiented head, wheeas those with positive values ae moe posteioly oiented. b: incipal component 2. Negative values indicate a shote distance between the adial tubeosity and the 80% level of the shaft and a moe medially located styloid pocess, and positive values have a longe neck and moe anteioly located styloid pocess. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 158

190 Summay Lateal cuvatue is the most impotant C fo both the medial and the lateal shaft sufaces (mcuveamhc1 and lcuveamhc1). This is eflected in the significant coelation between the scoes fo the cuvatue Cs (=0.271) (Table 4-53). Thee is no coelation between the Cs of the epiphyses and the two cuvatue Cs (Table 4-54). Table 4-53 eason s coelation matix: adial cuvatue Cs (n= 360) mcuamhc1 lcuamhc <0.001** mcuveamhc ** mcuveamhc lcuvamhc <0.001** lcuvamhc <0.001** ** Coelation is significant at the 0.01 level (2-tailed). * Coelation is significant at the 0.05 level (2-tailed). mcuamhc2 mcuamhc3 lcuvamhc * Table 4-54 eason s coelation matix: adial cuvatue and epiphyses Cs (n= 349). EpiAMHC1 EpiAMHC lcuvamhc mcuveamhc ** mcuveamhc ** lcuvamhc lcuvamhc ** ** Coelation is significant at the 0.01 level (2-tailed). * Coelation is significant at the 0.05 level (2-tailed). mcuveamhc1 Coelations between the othe shaft shape Cs indicate that individuals who have a highe degee of medial cuvatue (mcuveamhc1) have an apex of cuvatue at midshaft (mcuveamhc2), less medial expansion of the poximal inteosseous cest and the mediolateal diection of the distal end of the medial suface (lcuveamhc2) and a less 159

191 sinusoidal shaft (lcuveamhc3). A highe degee of lateal cuvatue (lcuveamhc1) and inceased sinusoidal shape (lcuveamhc3) is coelated with an inceased development of the poximal inteosseous cest and inceased medial pojection of the adial notch (mcuveamhc2). Coelations between the epiphysis Cs show that a moe posteioly pojected head esults in a moe developed poximal inteosseous cest, a moe developed adial notch (mcuveamhc2) and a moe sinusoidal shape (lcuveamhc3 but see lcuveamhc2). Coelation coefficients ae significant but low (see Left and Right diffeences) Ulna pincipal components explained osteio suface (pcuve) The fist fou Cs of the posteio cuve analysis explain 34.2%, 22.6%, 13.3% and 6.43%, espectively, of the vaiation (total 76.53%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. The distibution of the populations along C1 and 2 is shown in Figue C1 eflects diffeences in mediolateal cuvatue (Figue 4-52 and Figue 4-53a). C2 is the sinusoidal shape of the shaft in the mediolateal plane (Figue 4-52 and Figue 4-53b). C3 elates to the sinusoidal shape of the shaft in the anteoposteio plane (Figue 4-53c) and best eflects the posteio subtense descibed in the liteatue (Fische, 1904). C4 is the diection of the poximal shaft (Figue 4-53d). 160

192 Figue 4-52 The fist and second Cs fo the posteio cuve of the ulna. All ecent moden human samples. Cs ae explained in Figue

193 a b c d Figue 4-53 Mophological tends fo the posteio cuvatue of the ulna fo all ecent moden humans. a: incipal component 1: anteio view. Negative values have a highe degee of mediolateal cuvatue, wheeas positive values have a lowe degee of cuvatue. b: incipal component 2: anteio view. ositive values have a staight shaft while negative values ae sinusoidal in the mediolateal plane. c: incipal component 3: medial view ositive values ae moe sinusoidal in the anteoposteio plane compaed to negative values. d: incipal component 4: medial view. ositive values show a bent poximal shaft indicating a moe anteioly pojected ulna head, wheeas negative values ae elatively staight. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 162

194 oximal ulna (pox) The fist fou Cs of the poximal ulna analysis explain 22.0%, 18.4%, 7.84% and 4.33%, espectively, of the vaiation (total 52.6%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. C1 eflects diffeences in the oientation of the poximal ulna in elation to the shaft (Figue 4-54 and Figue 4-55a). C2 elates to the distance between the 80% level of the shaft and the coonoid pocess (Figue 4-55b). C3 shows the oientation of the tochlea notch (Figue 4-55c). C4 is elated to the size and dimensions of the tochlea notch (Figue 4-55d). opulation distibution fo the oientation of the poximal ulna and the distance between the 80% level of the shaft and the coonoid pocess is shown in Figue Figue 4-54 The fist and second Cs fo the poximal ulna. All ecent moden human samples. Cs ae explained in figue Figue

195 a b c d Figue 4-55 Mophological tends fo the poximal ulna fo all ecent moden humans. a: incipal component 1. Negative values have a poximal ulna that is medially pojected with a medial facing tochlea notch, wheeas positive values have a head that is lateally pojected and has a moe lateal facing tochlea notch. b: incipal component 2. ositive values have a longe distance between the 80% and the coonoid pocess, wheeas negative values have shot distances. C3 shows the oientation of the tochlea notch. c: incipal component 3. Negative values have a moe poximo-anteio facing tochlea notch and positive values have a moe anteio facing tochlea notch. d: incipal component. ositive values have a deepe tochlea notch with a highe adial notch and a lowe olecanon pocess compaed to the negative values. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 164

196 Summay Because the analysis of the ulna has the main goal of identifying the coelates with adial cuvatue, and the bone is one that is not fequently studied, the analyses below ae exploatoy and will conside all the Cs descibed above. The coelations between the posteio cuve and the poximal ulna Cs shows thee is a negative coelation between the distance between the 80% level of the shaft and the coonoid pocess (poxamhc2) and the sinusoidal shape in the anteoposteio plane. Individuals with a geate distance between the 80% level of the shaft and the coonoid pocess have a moe sinusoidal shaft shape in the anteoposteio plane (Table 4-55). Table 4-55 eason s coelation matix: posteio suface and poximal ulna Cs (n= 347). poxamhc1 poxamhc * pcuveamhc pcuveamhc ** <0.001 pcuveamhc * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). pcuveamhc poxamhc poxamhc Coelations between Cs and univaiate measuements All moden humans Cuvatue of the medial cuve of the adius is positively coelated with obusticity of the head and distal aticulation. McuveAMHC2 and lcuveamhc3 ae negatively coelated with obusticity of the distal aticulation and show that individuals with elatively lage distal aticulations have a medial pojection on the poximal inteosseous cest and a moe ponounced ulna notch (mcuveamhc2) and ae moe sinusoidal (lcuveamhc3) compaed to those with smalle distal aticulations. Thee is also a positive coelation between midshaft and head 165

197 obusticity and a moe sinusoidal adius (N=35) (Table 4-56). Lateal cuvatue of the adial shaft is not coelated with any of the univaiate measuements. Individuals with a elatively longe adial neck and an anteoposteioly naow head have a highe apex of cuvatue and a moe lateally pojecting styloid pocess (lcuveamhc2). Thee is a elationship between inceased obusticity of the distal aticulation, a elatively longe adial neck and a moe anteioly located styloid pocess (EpiAMHC2) (Table 4-56). Anteoposteio sinusoidal shape of the ulna shaft (pcuveamh3) is elated to the olecanon oientation, elative size of the poximal ulna and elative position of the bachial tubeosity. The mediolateal oientation of the poximal ulna (poxamhc1) is positively coelated with the coonoid-olecanon size atio, the size of the bachial tubeosity, length of the ponato cest and midshaft obusticity. Individuals with a shote distance between the tip of the coonoid pocess and the 80% level of the shaft (poxamhc2) have a smalle poximal ulna size, a smalle adial notch suface aea, a highe coonoid-olecanon size atio, a lage bachial tubeosity and inceased obusticity at the 25% level of the shaft and geate distal aticulation obusticity. Individuals with a moe poximoanteioly facing athe than an anteioly facing tochlea notch (poxamhc3) have a elatively smalle olecanon, a moe poximoanteioly facing tochlea notch, geate angle of the poximal ulna and inceased distal aticulation obusticity. The depth of the tochlea notch and the position of the adial notch (poxamhc4) ae positively coelated with the midshaft shape atio, the position of the adial notch and obusticity at the 25% of the shaft (Table 4-57; Table 4-58). 166

198 Table 4-56 eason s coelation matix fo adius shape Cs and univaiate measuements fo all moden human populations (N=35). mcuveamh C1 Midshaftobusticit y lcuvamh C Headobusticity ** distatshaftsizer 0.001** atio Max_ Length neck-shaft angle * osradtubml DosalST LatealST * NeckLengthRatio HeadShapeRatio midshaftshaperat 0.047* io * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). mcuveamh C * mcuveamh C * lcuvamh C ** ** lcuvamh C * ** ** ** <0.001 EpiAMH C * EpiAMH C *

199 Table 4-57 eason s coelation matix fo ulna shape Cs and univaiate measuements fo all moden human populations (N=35). Max_ Length Olec-shaftatio MidShaftShape pcuveamhc pcuveamhc pcuveamhc * pcuveamhc oxamhc oxamhc ** oxamhc * oxamhc ** * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). Radial Notch Suface atio ** < TochNotchOi ** Olec-oient angle * ** < * *

200 Table 4-58 eason s coelation matix fo ulna shape Cs and univaiate measuements fo all moden human populations (N=35). CoOleRatio BachRatio pon.c. length pcuveamhc pcuveamhc * pcuveamhc oxamhc ** ** oxamhc ** ** oxamhc oxamhc * * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). pcuveamhc1 Robust 50% ** < Robust 25% ** ** Robust dist at ** < *

201 opulations with high activity levels only Fo high activity goups, lateal cuvatue of the adius is not coelated with obusticity. Medial cuvatue is coelated with the obusticity of the aticulations, but not with obusticity at midshaft. Robusticity of the distal aticulation is negatively coelated with mcuveamhc2 and lcuveamhc3 and shows that individuals with elatively lage distal aticulations have an inceased medial extension of the poximal inteosseous cest, a medial diection of the distal cuve (moe medially expanded ulna notch) and ae moe sinusoidal compaed to those with smalle distal aticulations (Table 4-59). The coelation between the shape Cs of the ulna and the univaiate measuements ae the same as fo the whole sample. Only, thee is no elationship between the depth of the tochlea notch (oxamhc4) and the position of the adial notch and obusticity at the 25% of the shaft (Table 4-66; Table 4-67) 170

202 Table 4-59 eason s coelation matix fo adius Cs and univaiate measuements fo populations with high activity levels (N=20). mcuvea lcuvamh mcuveamh MHC1 C1 C2 Midshaftobusticity Headobusticity * distatshaftsizeratio ** * Max_ Length neck-shaft angle osradtubml DosalST LatealST NeckLengthRatio HeadShapeRatio midshaftshaperatio * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). mcuveamh C * lcuvamh C ** * lcuvamh C * * <0.001** EpiAMHC ** EpiAMHC

203 Table 4-60 eason s coelation matix fo ulna Cs and univaiate measuements fo populations with high activity levels (N=19) Max_ Length Olec-shaftatio MidShaftShape pcuveamhc pcuveamhc pcuveamhc * pcuveamhc oxamhc oxamhc * oxamhc * oxamhc * * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). Rad.Notch Suf ** TochNotchOi ** Olec-oient angle * ** < *

204 Table 4-61: eason s coelation matix fo ulna Cs and univaiate measuements fo populations with high activity levels (N=19). CoOleRatio BachRatio pon.c. length pcuveamhc pcuveamhc pcuveamhc ** pcuveamhc oxamhc ** 0.502* 0.737** <0.001 oxamhc * ** oxamhc oxamhc * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). Robusticity at 50% ** Robusticity at 25% * * Robust dist atic ** < **

205 Factos influencing cuvatue in moden humans The following analyses will focus on the elationship between adial cuvatue and ulna shaft shape and the behavioual, envionmental and biological vaiables that might be expected to influence mophology. These analyses addess the same hypotheses and pedictions tested fo the femu Bilateal asymmety of the lowe am Left and ight side ae not diffeent in degee of adial cuvatue (Table 4-71) (N=143 left and 218 ight adii). Left adii have a moe developed poximal inteosseous cest and adial notch (mcuveamhc2) and a staighte shaft, wheeas the ight adius is moe sinusoidal and lacks the poximal development on the inteosseous cest (mcuveamhc3) (Table 4-71). Left adii have a moe posteioly oiented adial head (EpiAMHC1) than ight adii. The high t-value fo EpiAMHC1 indicates that the shape diffeences along the C axis tanslate into the diffeences between ight and left (Table 4-62). The ulna shows maked asymmety. Right ulnae have moe medial cuvatue (pcuveamhc1) and ae moe sinusoidal in the mediolateal plane than left ulnae (pcuveamhc2) (Table 4-63). Right ulnae have a poximal ulna that is medially pojected with a medial facing tochlea notch (poxamhc1), have a moe poximo-anteio tochlea notch (poxamhc3), and a deepe tochlea notch with a highe adial notch and a lowe olecanon pocess (poxamhc4) (Table 4-63). 174

206 Table 4-62 Student s t-test esults fo bilateal asymmety in adius shape in moden humans. Side left ight left ight left ight left ight left ight left ight left ight left ight McuveAMHC1 lcuvamhc1 McuveAMHC2 McuveAMHC3 lcuvamhc2 lcuvamhc3 EpiAMHC1 EpiAMHC2 N Mean S.D t <0.001** <0.001** <0.001** * Significant at α=0.05 Table 4-63 Student s t-test esults fo bilateal asymmety in ulna shape in moden humans. cuveamhc1 cuveamhc2 cuveamhc3 cuveamhc4 poxamhc1 poxamhc2 poxamhc3 poxamhc4 Side ight left ight left ight left ight left ight left ight left ight left ight left N Mean S.D t * * <0.001* * <0.001* *=significant at α=0.05 Univaiate measuements Left adii have lowe neck-shaft angles, a moe medially located adial tubeosity, and a highe dosal and lateal subtense (Table 4-64). Right ulnae have lage poximal ulnae (Olec-shaftatio) that ae oiented moe in line with the shaft axis both mediolateally (head oientation angle) and anteoposteioly (toch-notch oientation) (Table 4-65). Right ulnae also have moe equal coonoid and olecanon heights 175

207 (CoOleRatio), a shote ponato cest and lowe obusticity at midshaft and at the 25% level of the shaft. Table 4-64 Student s t-test esults fo univaiate measuements of the adius in moden humans. Max_ Length neck-shaft angle osradtubml DosalST LatealST NeckLengthRatio OlecShapeRatio Midshaft Shape Ratio left ight left ight left ight left ight left ight left ight left ight left ight N Mean S.D t * * * * * Significant at α=0.05 Table 4-65 Student s t-test esults fo univaiate measuements of the ulna in moden humans. Max_ Length Olec-shaft atio MidShaft Shape Radial Notch Suf. atio TochNotchOi Olec-oient angle CoOleRatio BachRatio Rel. pon. c. size Robusticity at 50% Robusticity at 25% Robust dist atic Side ight left ight left ight left ight left ight left ight left ight left ight left ight left ight left ight left ight left N Mean S.D t * * * * <0.001* <0.001* * *=significant at α=

208 Results below ae epoted fo the pooled sample only, unless the significance values ae affected. In the analyses investigating sex diffeences all vaiables affected by bilateal asymmety ae pefomed fo the ight side only Body size Anteoposteio diamete of the femoal head is used as an measue of body size (fo those specimens fo which the femu is also peseved) (Ruff, 1991; McHeny, 1992; Gine et al., 1995). Based on this size suogate thee is no coelation between cuvatue of the adius and the shape of the ulna shaft and body size. (Table 4-66; Table 4-67). Table 4-66 eason s coelations fo body size (head diamete) and adial cuvatue (N=27). HeadAdiam mcuveamhc lcuvamhc * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). Table 4-67 eason s coelations fo body size (head diamete) and ulna shaft shape (N=27). HeadAdiam UlnpcuveAMHC UlnpcuveAMHC * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed) Sex As fo the femu the pupose of these analyses is to investigate sexual dimophism in the lowe am. Cuvatue Fo the whole sample of adii of known sex (N=90 males and 82 females), the pediction that 177

209 males have highe obusticity (Table 4-68) because they have highe activity levels than females was met but males and females wee not diffeent in degee of adial cuvatue (Table 4-69). Table 4-68 Student s t-test esults fo obusticity in moden human males and females. Midshaftobusticity Headobusticity distatshaftsizeratio Sex male female male female male female N Mean S.D t * * * * Significant at α=0.05 Table 4-69 Student s t-test esults fo adius cuvatue Cs in moden human males and females. mcuveamhc1 lcuvamhc1 Sex male female male female N Mean S.D t * Significant at α=0.05 It was demonstated in the analyses of the femu that thee is evidence that division of labou is most ponounced in goups with high activity levels. Theefoe, the expectation is that the effect of sex on obusticity and cuvatue is moe evident in those goups than fo the whole sample (Table 4-70; Table 4-71). The pediction is met fo obusticity (N=39 males and 38 females) and as is the case fo the whole sample, males and females ae significantly diffeent fo shaft and epiphyseal obusticity. Degee of cuvatue is not diffeent between males and females with high activity levels. Table 4-70 Student s t-test esults fo adius obusticity in moden human males and females with high activity levels. Sex N Mean S.D. t Midshaftobusticity male * female Headobusticity male <0.001* female distatshaftsizeratio male <0.001* female * Significant at α=

210 Table 4-71 Student s t-test esults fo adius cuvatue Cs in moden human males and females with high activity levels. mcuveamhc1 lcuvamhc1 Sex male female male female N Mean S.D t * Significant at α=0.05 Othe shaft shape Cs The othe shaft shape Cs ae bilateally asymmetic so only the ight side is analysed. Fo all individuals (N=61 males and 50 females) females have a moe ponounced poximal inteosseous cest and ulna notch (mcuveamhc2) and have a moe sinusoidal anteoposteio shaft shape (mcuveamhc3) (Table 4-72). The diffeence in anteoposteio shaft shape (mcuveamhc3) is also pesent in the goups with high activity levels, but not the diffeence in the inteosseous cest and the ulna notch (Table 4-73). Table 4-72 Student s t-test esults fo adius shaft shape Cs in moden human males and females ight only. mcuveamhc2 mcuveamhc3 lcuvamhc2 lcuvamhc3 Sex male female male female male female male female N Mean S.D t * * * Significant at α=

211 Table 4-73 Student s t-test esults fo adius shaft shape Cs in moden human males and females with high activity levels ight only. mcuveamhc2 mcuveamhc3 lcuvamhc2 lcuvamhc3 Sex male female male female male female male female N Mean S.D t * * Significant at α=0.05 The fist and second C fo the ulna show bilateal asymmety. Theefoe the analyses ae pefomed on the ight side only. Thee is no sexual dimophism in the ight ulna (65 males and 49 females) fo any of the shaft shape Cs (Table 4-74). In the ight ulnae of the high activity goup (30 males and 22 females), males have a staighte shaft in the mediolateal plane compaed to females (pcuveamhc2) (Table 4-75). Table 4-74 Student s t-test esults fo ulna shaft shape Cs in moden human males and females ight only. pcuveamhc1 pcuveamhc2 pcuveamhc3 pcuveamhc4 Sex male female male female male female male female N Mean S.D t *=significant at α=0.05 Table 4-75 Student s t-test esults fo ulna shaft shape Cs in moden human males and females with high activity levels ight only. pcuveamhc1 pcuveamhc2 pcuveamhc3 pcuveamhc4 Sex male female male female male female male female N Mean S.D t * *=significant at α=

212 Epiphysis shape EpiAMHC1 shows significant side diffeences so only the ight side is analysed. Males and females ae simila in thei adial epiphysis mophology fo the whole sample (Table 4-76) and fo high activity goups alone (Table 4-77). Table 4-76 Student s t-test esults fo adius epiphyses Cs in moden human males and females ight only. EpiAMHC1 EpiAMHC2 Sex male female male female N Mean S.D t * Significant at α=0.05 Table 4-77: Student s t-test esults fo adius epiphyses Cs in moden human males and females with high activity levels ight only. EpiAMHC1 EpiAMHC2 Sex male female male female N Mean S.D t * Significant at α=0.05 Thee is bilateal asymmety in the Cs fo the poximal ulna. Theefoe, these analyses ae pefomed on the ight ulna only. Males have a longe distance between the 80% level of the shaft and the tip of the coonoid pocess than females (65 males and 49 females) (poxamhc2) (Table 4-78). Fo ight ulnae of the high activity goups (30 males and 22 females) thee is no diffeence in poximal ulna shape (Table 4-79). Table 4-78 Student s t-test esults fo the poximal ulna Cs in moden human males and females ight only. poxamhc1 poxamhc2 poxamhc3 poxamhc4 Sex male female male female male female male female N Mean S.D t * *=significant at α=

213 Table 4-79 Student s t-test esults fo the poximal ulna Cs in moden human males and females with high activity levels ight only. poxamhc1 poxamhc2 poxamhc3 poxamhc4 Sex male female male female male female male female N Mean S.D t *=significant at α=0.05 Univaiate measuements Males have significantly longe adii (Max_Length, p<0.001 fo both all AMH and high activity goups only) (Table 4-80). When all individuals ae consideed females have a elatively shote adial neck (NeckLengthRatio), but this sexual dimophism disappeas when only goups with high activity levels ae consideed (Table 4-81). Table 4-80: Student s t-test esults fo the univaiate measuements of the adius in moden human males and females. Undelined vaiables show bilateal asymmety and wee analysed fo the ight side only. Max_ Length neck-shaft angle osradtubml DosalST LatealST NeckLengthRatio HeadShapeRatio midshaftshaperation Sex male female male female male female male female male female male female male female male female N Mean S.D t <0.001* * * Significant at α=

214 Table 4-81 Student s t-test esults fo the univaiate measuements of the adius in moden human males and females with high activity levels. Undelined vaiables show bilateal asymmety and wee analysed fo the ight side only. Max_ Length neck-shaft angle osradtubml DosalST LatealST NeckLengthRatio HeadShapeRatio midshaftshaperation Sex male female male female male female male female male female male female male female male female N Mean S.D t <0.001* * Significant at α=0.05 Males have longe ulnae than females (Max_Length, p<0.001 fo both all AMH and high activity goups only) (Table 4-82). When all individuals ae consideed, males ae ounde at the ulna midshaft and ae moe obust at the midshaft and at the 25% level of the shaft than females. These diffeences disappea when only high activity goups ae analysed (Table 4-83). 183

215 Table 4-82 Student s t-test esults fo the univaiate measuements of the ulna in moden human males and females. Undelined vaiables show bilateal asymmety and wee analysed fo the ight side only. Max_ Length Olecshaftation MidShaftShape Radial Notch Suface atio TochNotchOi Olec-oient angle CoOleRation bachratio Rel. pon. c. size Robusticity at 50% Robusticity at 25% Robust dist atic Sex male female male female male female male female male female male female male female male female male female male female male female male female N Mean S.D t <0.001* * * < * * * *=significant at α=

216 Table 4-83 Student s t-test esults fo the univaiate measuements of the ulna in moden human males and females with high activity levels. Undelined vaiables show bilateal asymmety and wee analysed fo the ight side only. Max_ Length Olec-shaftatio MidShaftShape Radial Notch Suface atio TochNotchOi Olec-oient angle CoOleRation bachratio Size pon.c. el. length Robusticity at 50% Robusticity at 25% Robust dist atic Sex male female male female male female male female male female male female male female male female male female male female male female male female N Mean S.D t <0.001* *=significant at α=0.05 Summay Thee is no sexual dimophism in cuvatue of the shaft of the adius o the ulna. Females have shote and less obust adii than do males. Females also have a moe anteoposteio sinusoidal adial shaft shape. Females have a smalle ulna that is moe sinusoidal than that of males Age Of the whole sample, thee wee 93 adii fom individuals of known age and 97 ulnae. Thee is no elationship between age and cuvatue and epiphyseal shape of the adius (Table 4-84), fo the pooled and ight-only sample. A negative coelation with mcuveamhc2 and lcuveamhc3 indicate that younge individuals have an inceased medial extension of the 185

217 poximal inteosseous cest and a medial diection of the distal cuve (moe medially expanded ulna notch) and have a moe sinusoidal shape compaed to olde individuals. Table 4-84 Kendall s Tau b coelations fo adius Cs and age (N=93). Specimen age Cuvatue McuAllC1 LcuAllC1 Shaft shape McuAllC2 McuAllC3 LcuAllC2 LcuAllC ** ** <0.001 Epiphyses shape EpiAllC EpiAllC ** <0.001 * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). Thee is no elationship between age and the shape of the shaft o the poximal ulna (Table 4-85). Thee is a weak coelation (=0.182, =0.041) between poxamhc1 and age fo the ight ulnae. Olde individuals have a moe medially pojected poximal ulna with a moe medial facing tochlea notch. Table 4-85 Kendall s Tau b coelations fo ulna Cs (N=97). shaft shape pcuveamhc1 oximal ulna poxamhc pcuveamhc poxamhc pcuveamhc poxamhc pcuveamhc poxamhc * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed)

218 Univaiate measuements Age is positively coelated with head-shaft size atio (Table 4-86). Olde individuals have lage heads elative to shaft length compaed to younge individuals. Fo the ulna the position of the bachial tubeosity may shift moe distally but this coelation is absent in the ight only sample (=0.163; =0.068). Thee is a weak positive coelation between distal aticulation obusticity and age (Table 4-87). Table 4-86 Kendall s Tau b coelations fo the univaiate measuements on the adius and age (N=97). Max_ Length neck-shaft angle osradtubml DosalST LatealST NeckLengthRatio HeadShapeRatio midshaftshaperatio HeadShaftSizeRatio Specimen age ** ** * Robusticity Midshaftobusticity Headobusticity 0.174* distatshaftsizeratio * = Coelation is significant at the 0.05 level (2tailed). ** = Coelation is significant at the 0.01 level (2tailed). 187

219 Table 4-87 Kendall s Tau b coelations fo univaiate measuements on the ulna (N=97). oximal ulna Olec-oient angle Olec-shaftatio 0.171* CoOleRatio MidShaftShape bachratio Rad. Notch Suf. atio Rel. pon.c. size TochNotchOi * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). Max_ Length ** obusticity Robusticity at 50% Robusticity at 25% Robust dist atic **

220 When age categoies wee used in ode test fo the effect of age on adial cuvatue the ANOVA showed no significant effect (Table 4-88). Fo the ulna, adults have the shotest distance between the 80% level of the shaft and the tip of the coonoid pocess (poxamhc2.) (Table 4-89, Figue 4-56). Table 4-88 ANOVA esults fo age categoies and adius cuvatue Cs. d.f.=2 F mcuveamhc lcuvamhc *=significant at α=0.05 Sig Table 4-89 ANOVA esults fo age categoies and ulna shape Cs. d.f.=2 pcuveamhc1 pcuveamhc2 pcuveamhc3 pcuveamhc4 poxamhc1 poxamhc2 poxamhc3 poxamhc4 *=significant at α=0.05 F Sig *

221 Figue 4-56 oxamhc2 fo moden humans, by age stategy. Mean and 95% confidence inteval (whiskes). High values have a geate distance between the tip of the coonoid pocess and the 80% level of the shaft. Summay Thee is no elationship between age and cuvatue, no ae thee cuvatue diffeences when the age categoies ae used. Younge individuals have an inceased medial extension of the poximal inteosseous cest and a medial diection of the distal cuve (moe medially expanded ulna notch) and have a moe sinusoidal shape compaed to olde individuals. Olde individuals have lage heads elative to shaft length compaed to younge individuals Activity levels The following analyses use the same categoies used in the analyses of the femu. The distibution of the populations (Appendix 8) fo the fist Cs of the adius and ulna ae pesented in Figue 4-57 to Figue 4-59). 190

222 Figue 4-57 Distibution of the activity level categoies in the space of C1 (degee of cuvatue) and C2 (medial expansion of the inteosseous cest) of the medial cuve of the adius fo all moden humans. Cicles: high activity, squaes: modeate activity, cosses: low activity. 191

223 Figue 4-58 Distibution of the activity level categoies in the space of C1 (degee of cuvatue) and C2 (apex of cuvatue) of the lateal cuve of the adius fo all moden humans. Cicles: high activity, squaes: modeate activity, cosses: low activity. 192

224 Figue 4-59 Distibution of the activity level categoies in the space of the C1 (degee of mediolateal cuvatue) and C2 (mediolateal sinusoidal shape) of the posteio cuve of the ulna fo all moden humans. Cicles: high activity, squaes: modeate activity, cosses: low activity. Cuvatue Thee ae no diffeences in the cuvatue of the adius acoss activity levels(table 4-90). Thee ae diffeences between high activity subsistence stategies fo both cuvatue Cs, howeve (Table 4-91). The hoticultualists ae the least cuved and significantly diffeent in lateal cuvatue fom equestian foages and pastoalists (lcuveamhc1) (Appendix 23,Figue 4-60). The post-hoc compaisons show no significant paiwise diffeences fo mcuveamhc1. Table 4-90 ANOVA esults fo activity levels and adius cuvatue Cs. d.f.=2 F mcuveamhc lcuvamhc *=significant at α=0.05 Sig

225 Table 4-91 ANOVA esults fo high activity subsistence subsistence stategies and adius cuvatue Cs. d.f.=4 mcuveamhc1 lcuvamhc1 *=significant at α=0.05 F Sig * 0.002* Figue 4-60 Lateal cuvatue of the adius fo moden humans, by subsistence stategy. The scale of lcuveamhc1 is evesed so that the highe values indicate a highe degee of cuvatue. Mean and 95% confidence inteval (whiskes). Othe shaft shape Cs The diffeent activity level goups ae significantly diffeent in adial shaft shape in one out of fou Cs (lcuveamhc3) (Table 4-92). ost-hoc tests of the lcuveamhc3 show that high activity goups have the staightest shaft compaed to the moe sinusoidal shaft in modeate and low activity goups (Appendix 24 and Figue 4-61). 194

226 Table 4-92 ANOVA esults fo activity levels and adius shaft shape Cs. d.f.=2 mcuveamhc2 mcuveamhc3 lcuvamhc2 lcuvamhc3 *=significant at α=0.05 F Sig * Figue 4-61 LcuvAMHC3 (high values have the least sinusoidal shaft) of the adius fo moden humans, by high activity subsistence stategy. Mean and 95% confidence inteval (whiskes). Fo two out of fou othe adial shaft shape Cs the high activity subsistence categoies ae significantly diffeent (Table 4-93). Compaed to pedestian foages, aquatic foages have an inceased medial extension of the poximal inteosseous cest and a medial diection of the distal cuve (moe medially expanded ulna notch) (mcuveamhc2) (Figue 4-62). astoalists have the most sinusoidal shaft compaed to othe subsistence categoies (lcuveamhc3) (Appendix 25) (Figue 4-63). Fo the ulna, thee ae no significant diffeences in shaft shape between the diffeent activity goups o subsistence pattens (Table 4-94; Table 4-95). 195

227 Table 4-93 ANOVA esults fo subsistence stategies with high activity levels and adius shaft shape Cs. d.f.=4 F mcuveamhc mcuveamhc lcuvamhc lcuvamhc *=significant at α=0.05 Sig * * Figue 4-62 McuveC2 of the adius fo moden humans, by high activity subsistence stategy. Mean and 95% confidence inteval (whiskes). Low values have inceased medial extension of the poximal inteosseous cest. 196

228 Figue 4-63 LcuveC3 (low values ae moe sinusoidal) of the adius fo moden humans, by high activity subsistence stategy. Mean and 95% confidence inteval (whiskes). Table 4-94 ANOVA esults fo activity levels and ulna shaft shape Cs. d.f.=2 pcuveamhc1 pcuveamhc2 pcuveamhc3 pcuveamhc4 *=significant at α=0.05 F Sig Table 4-95 ANOVA esults fo subsistence stategies with high activity levels and ulna shaft shape Cs. d.f.=4 pcuveamhc1 pcuveamhc2 pcuveamhc3 pcuveamhc4 *=significant at α=0.05 F Sig

229 Epiphysis shape Although thee is a significant diffeence between the activity levels fo adial epiphysis shape, the post-hoc pocedues did not find diffeences between the thee activity levels (Table 4-96 and Appendix 26). Fo the high activity subsistence goups thee is a significant diffeence between pastoalists and aquatic and equestian foages fo EpiAMHC2 (Table 4-97). astoalists have a moe posteioly oiented head than hoticultualists and aquatic and equestian foages (Figue 4-64 and Appendix 27). Table 4-96 ANOVA esults fo activity levels and adius epiphyses Cs. d.f.=2 EpiAMHC1 EpiAMHC2 *=significant at α=0.05 F Sig * Table 4-97 ANOVA esults fo subsistence goups with high activity levels and adius epiphyses Cs. d.f.=4 EpiAMHC1 EpiAMHC2 *=significant at α=0.05 F Sig. <0.001*

230 Figue 4-64 EpiAMHC2 fo moden humans, by subsistence stategy. The scale of lcuveamhc1 is evesed to ease intepetations (have a moe posteioly oiented head). Mean and 95% confidence inteval (whiskes). High values have a moe posteioly oiented head. The activity levels ae significantly diffeent fo poxamhc4 (Table 4-98). opulations with low activity levels have a deepe tochlea notch with a highe adial notch and a lowe olecanon pocess compaed to the high and modeate activity goups (Appendix 27, Figue 4-65). 199

231 Table 4-98 ANOVA esults fo activity levels and the poximal ulna Cs. d.f.=2 poxamhc1 poxamhc2 poxamhc3 poxamhc4 *=significant at α=0.05 F Sig <0.001* Figue 4-65 oxamhc4 fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). High values have a deepe tochlea notch with a highe adial notch and a lowe olecanon pocess. The high activity subsistence goups ae diffeent fo poxamhc2 and poxamhc4 (Table 4-99). Equestian and aquatic foages have a geate distance between the 80% shaft level and the tip of the coonoid pocess compaed to pastoalists who have the shotest distance. Also, aquatic foages have a shallowe tochlea notch with a lowe adial notch and a highe olecanon pocess compaed to pastoalists and pedestian and equestian foages (Figue 4-66 and Figue 4-67) (Appendix 29). 200

232 Table 4-99 ANOVA esults fo subsistence stategy and the poximal ulna Cs. d.f.=4 poxamhc1 poxamhc2 poxamhc3 poxamhc4 *=significant at α=0.05 F Sig * * 201

233 Figue 4-66 oxamhc2 fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). High values geate distance between the tip of the coonoid pocess and the 80% level of the shaft. Figue 4-67 oxamhc4 fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). High values have a deepe tochlea notch with a highe adial notch and a lowe olecanon pocess. 202

234 Univaiate measuements The activity goups ae diffeent fo adial head obusticity (Table 4-100; Appendix 30; Figue 4-68). The high activity subsistence stategies ae significantly diffeent in obusticity at all thee levels of the adial shaft (head, midshaft and distal aticulation) (Table 4-101). astoalists ae the most obust oveall, wheeas hoticultualists and aquatic foages have the least obust adii (Appendix 31 and Figue Figue 4-71). Table ANOVA esults fo activity level and adius obusticity. d.f.=2 Midshaftobusticity Headobusticity distatshaftsizeratio *=significant at α=0.05 F Sig <0.001* Figue 4-68 Head obusticity fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). 203

235 Table ANOVA esults fo subsistence stategy and adius obusticity. d.f.=4 F Midshaftobusticity Headobusticity distatshaftsizeratio *=significant at α=0.05 Sig * 0.001* 0.001* Figue 4-69 Midshaft obusticity fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). 204

236 Figue 4-70 Head obusticity fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). Figue 4-71 Relative distal aticulation size fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). 205

237 Thee is a significant diffeence fo the activity goups fo a seies of univaiate measuments of the adius. Low activity goups have a smalle neck-shaft angle and a moe medially placed adial tubeosity (Table 4-102; Figue 4-72 and Figue 4-73). High activity goups have a shote neck than both modeate and low activity goups and less dosal subtense than modeate activity goups (Appendix 31 and Figue 4-74 and Figue 4-75). The diffeences fo two univaiate measuements affected by bilateal asymmety disappea when only the ight side is consideed (Neck-shaft angle and position and adial tubeosity) (Appendix 33; Table 4-103). Table ANOVA esults fo activity level and univaiate measuements on the adius. d.f.=2 Max_ Length neck-shaft angle osradtubml DosalST LatealST NeckLengthRatio HeadShapeRatio midshaftshaperation *=significant at α=0.05 F Sig * 0.013* 0.031* <0.001* Table ANOVA esults fo activity level and univaiate measuements on the adius ight only. d.f.=2 Max_ Length neck-shaft angle osradtubml DosalST LatealST NeckLengthRatio HeadShapeRatio midshaftshaperatio *=significant at α=0.05 F Sig * <0.001*

238 Figue 4-72 osition of the adial tubeosity fo moden humans, by activity level. Lowe values ae moe medially placed. Mean and 95% confidence inteval (whiskes). Figue 4-73 Neck-shaft angle fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). 207

239 Figue 4-74 Relative adial neck length fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Figue 4-75 Dosal subtense fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). 208

240 Within the high activity goups (Table 4-104; Figue Figue 4-81) the aquatic foages have significantly shote adii with a high neck-shaft angle (Figue 4-76). Equestian foages have the lowest neck-shaft angle (Figue 4-77). astoalists have the lowest midshaft shape atio, indicating a moe developed inteosseous cest on the adius compaed to pedestian foages, hoticultualists and aquatic foages who have highe midshaft shape atios (Appendix 34; Figue 4-81). Table ANOVA esults fo subsistence stategy and univaiate measuements on the adius. d.f.=4 Max_ Length neck-shaft angle osradtubml DosalST LatealST NeckLengthRatio HeadShapeRatio midshaftshaperatio *=significant at α=0.05 F Sig. <0.001* <0.001* 0.036* 0.035* * <0.001* Figue 4-76 Maximum length fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). 209

241 Figue 4-77 Neck-shaft angle fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). Figue 4-78 osition of the adial tubeosity fo moden humans, by subsistence stategy. Lowe values ae moe medially placed. Mean and 95% confidence inteval (whiskes). 210

242 Figue 4-79 Dosal subtense fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). Figue 4-80 Neck length atio fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). 211

243 Figue 4-81 Midshaft shape atio fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). The activity goups ae significantly diffeent fo most univaiate measuements of the ulna (Table 4-105). High activity goups have shote ulnae, smalle adial notches, a smalle coonoid-olecanon size atio and ae moe obust than modeate activity goups (Appendix 35 and Figue Figue 4-90). Low activity goups ae moe obust at the 25% level of the shaft, have a lowe bachial tubeosity and a highe midshaft shape atio than do high and modeate activity goups (Figue 4-88 and Figue 4-83). 212

244 Table ANOVA esults fo activity levels and univaiate measuements on the ulna. d.f.=2 Max_ Length Olec-shaftatio MidShaftShape Rad. Notch Suf. atio TochNotchOi Olec-oient angle CoOleRatio bachratio Size pon.c. el. length Robusticity at 50% Robusticity at 25% Robust dist atic *=significant at α=0.05 F Sig * * 0.002* 0.004* 0.041* 0.009* 0.001* * 0.004* Figue 4-82 Maximum length fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). 213

245 Figue 4-83 Midshaft shape fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Figue 4-84 Radial notch suface aea fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). 214

246 Figue 4-85 Tochlea notch oientation fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Figue 4-86 Olecanon oientation angle fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). 215

247 Figue 4-87 Coonoid-olecanon atio fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Figue 4-88 Bachial muscle attachment atio fo moden humans, by activity level. Highe values have a elatively lowe insetion. Mean and 95% confidence inteval (whiskes). 216

248 Figue 4-89 Robusticity at 50% shaft level fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). Figue 4-90 Robusticity at 25% shaft level fo moden humans, by activity level. Mean and 95% confidence inteval (whiskes). 217

249 Thee ae significant diffeences between the high activity subsistence categoies fo all univaiate measuements of the ulna (except midshaft shape) (Table and Appendix 36). Aquatic foages have the shotest ulnae and have the elatively lagest poximal ulnae (Figue 4-91). The pastoalists have the lagest adial notch suface. The aquatic foages and the hoticultualists have the smallest adial notch suface and ae diffeent fom the pedestian foages and pastoalists (Figue 4-94). astoalists have the lagest olecanon angle compaed to pedestian and equestian foages (Figue 4-96). The coonoid-olecanon atio is lagest in pastoalists and smallest in equestian foages. Aquatic and pedestian foages and hoticultualists ae intemediate (Figue 4-97). astoalists have the lowest bachilis insetion, the hoticultualists the highest (Figue 4-92). The hoticultualists also have the longest ponato cest, wheeas equestian foages have the shotest (Figue 4-98). The equestian foages have the lowest midshaft obusticity, pastoalists the highest (Figue 4-99). At the 25% level of the shaft, pastoalists ae still the most obust, but the least obust ae the hoticultualists (Figue 4-100). Table ANOVA esults fo subsistence pattens and univaiate measuements on the ulna. d.f.=4 Max_ Length Olec-shaftatio MidShaftShape Radial Notch Suface atio TochNotchOi Olec-oient angle CoOleRation BachRatio Size ponato cest el. lenth Robusticity at 50% Robusticity at 25% Robust dist atic *=significant at α=0.05 F Sig. <0.001* 0.004* <0.001* 0.038* 0.013* <0.001* 0.042* 0.026* <0.001* <0.001* 0.009* 218

250 Figue 4-91 Ulna maximum length fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). Figue 4-92 osition of the bachial tubeosity fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). 219

251 Figue 4-93 Olecanon-shaft size atio fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). Figue 4-94 Radial notch suface atio fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). 220

252 Figue 4-95 Tochlea notch oientation fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). Figue 4-96 Olecanon oientation angle fo moden humans, by subsistence stategy. Mean and 95% confidence inteval (whiskes). 221

253 Figue 4-97 Coonoid-olecanon atio fo moden humans, by subsistence patten. Mean and 95% confidence inteval (whiskes). Figue 4-98 Relative size of the ponato cest fo moden humans, by subsistence patten. Mean and 95% confidence inteval (whiskes). 222

254 Figue 4-99 Robusticity at 50% shaft level fo moden humans, by subsistence patten. Mean and 95% confidence inteval (whiskes). Figue Robusticity at 25% shaft level fo moden humans, by subsistence patten. Mean and 95% confidence inteval (whiskes). 223

255 Figue Robusticity of the distal aticulation fo moden humans, by subsistence patten. Mean and 95% confidence inteval (whiskes) Evolution ove time in Euope Cuvatue Fo the adius, only the lateal suface cuvatue (lcuveamhc1) is significantly affected though time. Howeve; it doesn not show a steady decease.the Medieval populations ae the least lateally cuved (Figue 4-102) (Table 4-107) (Appendix 37; Appendix 8). Table ANOVA esults fo time-peiod and cuvatue of the adius. d.f.=3 mcuveamhc1 lcuvamhc1 *=significant at α=0.05 F Sig * 224

256 Figue Lateal cuvatue of the adius fo moden Euopeans, by time peiod. Mean and 95% confidence inteval (whiskes). Ulna shape The time peiods ae significantly diffeent fo two of the ulna Cs (Table 4-108; Appendix 38), but none of the significant vaiables shows a steady change though time. The Neolithic individuals have moe anteoposteioly sinusoidal shafts than the Medieval sample (pcuveamhc3) (Figue 4-103) and the 18th and 19th Centuy sample has a deepe tochlea notch (poxamhc4) (Figue 4-104). Table ANOVA esults fo time-peiod and ulna shape. d.f.=3 pcuveamhc1 pcuveamhc2 pcuveamhc3 pcuveamhc4 poxamhc1 poxamhc2 poxamhc3 poxamhc4 *=significant at α=0.05 F Sig * * 225

257 Figue cuveamhc3 (high values have a moe anteoposteioly sinusoisal shaft) of the adius fo moden Euopeans, by time peiod. Highe values have moe anteoposteioly sinusoisal shafts. Mean and 95% confidence inteval (whiskes). Figue oxamhc4 (high values have a deepe tochlea notch) of the adius fo moden Euopeans, by time peiod. Highe values have a deepe tochlea notch. Mean and 95% confidence inteval (whiskes). 226

258 Summay Although thee ae some diffeences among samples fom diffeent time peiods thee ae no geneal tends fo aspects of adius and ulna shape though time in Euope Climate and latitude As fo the femu analysis, latitude is used hee as a geneal poxy fo climate (Appendix 8). Individuals fom highe latitudes have a highe degee of lateal adial cuvatue than those fom lowe latitudes (LcuveAMHC1) (Table 4-109; Figue 4-105). Thee ae no coelations between the adial epiphysis shape Cs and latitude (Table 4-109). The othe shaft shape Cs show that individuals fom highe latitudes have an inceased medial extension of the poximal inteosseous cest with amoe medial expanded ulna notch (mcuveamhc2) (Figue 4-106) and a moe sinusoidal shape than those living in low latitudes (mcuveamhc3) (Table and Figue 4-107). Table eason s coelations fo cuvatue, apex of cuvatue, diaphyseal shape and epiphyses shape Cs and latitude (climate) on the adius (N=34). Latitude Cuvatue mcuveamhc1 lcuvamhc1 Epiphyses shape EpiAMHC1 * ** * Othe diaphyseal shape mcuveamhc2 mcuveamhc3 lcuvamhc2 lcuvamhc EpiAMHC Coelation is significant at the 0.05 level (2-tailed). Coelation is significant at the 0.01 level (2-tailed) ** *

259 Figue Lateal cuvatue of the adius (lcuveamhc1) and latitude fo ecent moden humans. Figue Medial expansion of the inteosseous cest (mcuveamhc2) and latitude fo ecent moden humans. 228

260 Figue Sinusoidal shape of the adius (mcuveamhc3) and latitude fo ecent moden humans. Individuals fom high latitudes have shote distances between the 80% level of the shaft and the tip of the coonoid pocess (Table 4-110) (poxamhc2). (poxamhc2) (Figue 4-108) and a moe poximo-anteio tochlea notch (poxamhc3) (Figue 4-109). Table eason s coelations fo cuvatue, apex of cuvatue, diaphyseal shape and epiphyses shape Cs and latitude (climate) on the ulna (N=32). Latitude Shaft shape pcuveamhc1 oximal ulna oxamhc pcuveamhc oxamhc pcuveamhc oxamhc pcuveamhc oxamhc * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed) ** < *

261 latitude Figue Distance between the 80% shaft level and the tip of the coonoid pocess (poxamhc2) and absolute latitude fo ecent moden humans. latitude Figue Oientation of the tochlea notch (poxamhc3) and absolute latitude fo ecent moden humans. 230

262 Univaiate measuements Head and distal aticulation obusticity of the adius is positively coelated with absolute latitude but midshaft obusticity is not (Table 4-111; Figue Figue 4-112). Thee is no elationship between the univaiate measuements of the adius and absolute latitude (Table 4-112). Latitude has a pevasive effect on ulna shape as epesented by the univaiate measuements. oximal ulna size, adial notch suface aea, tochlea notch oientation, olecanon oientation, coonoid-olecanon atio, bachial tubeosity length and distal aticulation obusticity ae positively coelated with latitude (Table 4-113, Figue Figue 4-118). Table eason s coelations fo adius obusticity (head, midshaft and distal aticulation) and latitude (climate) (N=34). Midshaftobusticity Headobusticity Latitude ** * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). distatshaftsizeratio ** Figue Radius midshaft obusticity and latitude fo ecent moden humans. 231

263 Figue Radius distal aticulation obusticity and latitude fo ecent moden humans. Figue Radius head obusticity and latitude fo ecent moden humans. 232

264 Table eason s coelations fo univaiate measuements on the adius and latitude (climate) (N=34). Absolute Latitude neck-shaft angle osradtubml DosalST LatealST NeckLengthRatio HeadShapeRatio midshaftshaperation * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2tailed). Max_ Length Table eason s coelations fo univaiate measuements on the ulna and latitude (climate) (N=31). Max_ Length TochNotchOi Olec-shaftatio 0.590** Olec-oient <0.001 MidShaftShape CoOleRation Rad Not Suf 0.476** BachRatio * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed) ** ** < * ** pon.c. size Robust 50% Robust 25% Robust dist at * <

265 latitude Figue Scatteplot fo olecanon shaft atio and latitude fo ecent moden humans. latitude Figue Radial notch suface aea and latitude fo ecent moden humans. 234

266 latitude Figue Oientation of the tochlea notch and latitude fo ecent moden humans. latitude Figue Olecanon oientation angle and latitude fo ecent moden humans. 235

267 latitude Figue osition of the bachial muscle insetion and latitude fo ecent moden humans. latitude Figue Distal aticulation obusticity of the ulna and latitude fo ecent moden humans. 236

268 Summay Individuals fom highe latitudes have a highe degee of lateal adial cuvatue and medial expansion of the inteosseous cest. Individuals fom highe latitudes also have a lage poximal ulna, lage adial notch suface aea, a moe poximo-anteioly facing tochlea notch, a moe lateal olecanon oientation, a highe coonoid-olecanon atio, a lowe elative bachial insetion and geate distal aticulation obusticity Mantel test The esults fo the Mantel test ae summaised in Table and Table Thee is a significant coelation between the lateal cuve of the adius and tempeatue which is consistent with the analysis of latitude. Thee is no coelation with cuvatue o the whole adius shape, altitude and aveage ainfall. Table Results of the Mantel tests pefomed fo envionmental distance matices adius. lateal cuvatue altitude diffeences ainfall diffeences tempeatue diffeences * = eason coelation coefficient medial cuvatue R whole adius shape * Randomisation tests with 5000 pemutations show significance values of p<0.05 fo all significant coelations between matices. Ulna shaft shape is not coelated with altitude, ainfall, tempeatue o geogaphic distance. The diection of the poximal ulna (poxc1) is coelated with altitude. The distance between the 80% level of the shaft and the tip of the coonoid pocess is coelated with ainfall and tempeatue. 237

269 Table Results of the Mantel tests pefomed fo envionmental distance matices - ulna altitude diffeences ainfall diffeences tempeatue diffeences = eason coelation coefficient pcuve pcuve2 R poximal ulna * poximal ulna * * whole bone Randomisation tests with 5000 pemutations show significance values of p<0.05 fo all significant coelations between matices. 238

270 Summay Oveall, thee is no asymmety between the between left and ight adial cuvatue (medial o lateal). Fo the ulna thee is some asymmety in the medial cuvatue and the mediolateal sinusoidal shape of the shaft. The pedictions that cuvatue of the adius and ulna would be elated to body size and activity levels wee not met. Thee is no sexual dimophism in adial cuvatue o in ulna shaft shape but males ae moe obust. Thee ae no geneal tends though time in Euope o with individual age. Cuvatue does not vay significantly between goups with diffeent activity levels. Within high activity goups, hoticultualists show the lowest degee of lateal cuvatue, and the equestian foages and pastoalists show the highest degee. astoalists ae the most obust in both ulna and adius. Thee is a positive coelation between latitude and lateal adius cuvatue. The Mantel test also showed coelations between colde tempeatue and moe ponounced cuvatue. Specimens with moe obust adii have less medial cuvatue. A minoity of the analyses pesented hee was exploatoy athe than pefomed to addess specific pedictions. The significant esults fom these analyses wee used to aid the intepetation of long bone cuvatue. Howeve, thee wee a few significant esults which did not fit any a pioi expectation and fo which the significance was close to Theefoe, it is likely that these occued because multiple tests wee conducted on the same data, and the Bonfeoni coection was not applied (see section 3.3). These esults include the staighte poximal posteio femoal diaphyses of males, wheeas those of females slope posteioly (pcuveamhc2, Student s t-test, p=0.031), and the moe medial pojection of the poximal ulna and moe medially facing tochlea notch of olde individuals (poxamhc1 Kendall tau b; =0.182, =0.041). Thee ae a couple of cases whee the significance value is low, but the esults did not follow the pedicted tend and cannot be functionally explained. These esults ae a moe anteoposteio sinusoidal ulna shaft in Neolithic populations (pcuveamhc3: ANOVA; F=3.326, =0.022) and a deepe tochlea notch in the 18th-19th centuy sample (poxamhc4: ANOVA; F=4.881, =0.003). 239

271 4.4. Systemic effects of cuvatue The coelations between cuvatue of the diffeent bones ae weak (Table 4-116) (N=27 populations). osteio femoal cuvatue and medial adial cuvatue ae coelated. The anteoposteio sinusoidal shape of the ulna (posteio subtense) is coelated with lateal adial cuvatue and anteio femoal cuvatue. Table eason s coelations fo cuvatue and apex of cuvatue Cs between the femu, adius and ulna (N=218). FemacuAMH FemcuvAMH C1 C Radmcuve AMHC * RadlcuvAM HC UlnpcuveA MHC UlnpcuveA MHC * UlnpcuveA MHC UlnpcuveA MHC4 * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). RadmcuveAMH C RadlcuvAMH C *

272 4.5. Discussion Thee diffeent hypotheses wee poposed to explain the vaiation in long bone cuvatue between moden human populations in Chapte 2. The esults of the foegoing analyses will be discussed in elation to the pedictions of these hypotheses. Hypothesis 1: A high degee of cuvatue is elated to body size. Long bone cuvatue in mammals is allometically scaled with body size (Biewene, 1983; Swatz, 1990). Biewene (1983) suggested that inceased cuvatue is a mechanism by which animals educe bone stesses because cuvatue esponds moe apidly to body size incease than does bone coss-sectional aea. Loading of the femoal diaphysis in humans is popotional to body size (Ruff, 2000b) and mophological featues, such as obusticity, ae also allometically elated to body size. On this basis, a elationship between cuvatue and body size is pedicted in the load-beaing femu. The elationship is expected to be somewhat diffeent in the am as the ulna and adius ae not weight-beaing bones and, theefoe, ae not axially loaded. Femu Fo this sample, the esults demonstate that thee is a elationship between extenal obusticity and body size (estimated using femoal head diamete as a poxy) fo both the femu and adius. Howeve, none of the cuvatue Cs of the femu ae coelated with body size, except in the populations with high activity levels whee division of labou is most ponounced. Lowe am Extenal obusticity is elated to body size (estimated using femoal head diamete). Thee is no coelation between the cuvatue of the adius and the ulna and body size. Thee is a diffeence between males and females in foeam obusticity but not in cuvatue of the adius. Females with high activity levels have a moe mediolateal sinusoidal shape compaed to males, but this diffeence is not pesent fo the whole sample. Although thee is a diffeence in femoal cuvatue between males and females in populations with high activity levels, the lack of sexual dimophism in long bone cuvatue fo the whole 241

273 sample suggests that the diffeences ae not due to the fact that males have lage bodies o because they have diffeent homone levels. Hypothesis 2: Cuvatue is a esponse to inceased activity levels. Femu Femoal cuvatue has two aspects. The fist is the degee of cuvatue and the second is the position of the apex of cuvatue. These ae not statistically covaiates but they behave similaly in thei elationships with habitual behaviou and envionmental factos. In geneal, as degee of cuvatue inceases, the apex of cuvatue moves infeioly. This confims the hypothesis suggested by Shackelfod and Tinkaus (2002) that a high degee of cuvatue is associated with a moe infeio apex of cuvatue. Individuals fom populations with high activity levels have moe cuved femoa and have a lowe apex of cuvatue than those fom populations with modeate and low activity levels. This elationship with activity is also eflected in a coelation of femoal cuvatue with skeletal measues of activity such as extenal obusticity. It was pedicted that if cuvatue and obusticity wee elated that thee would be a decease in femoal cuvatue occuing with agicultue and then with ubanism (Ruff et al., 1993; Tinkaus and Ruff, 1999b; Ruff and Tinkaus, 2000; Holt, 2003). This is confimed in the tempoal tend fo femoal cuvatue in the Euopean sample and suppots the hypothesis suggested by Shackelfod and Tinkaus (2002) that low levels of cuvatue ae elated to a decease in long-distance mobility. Thee is also no tend in cuvatue with inceasing age and deceasing activity intensity, howeve. Fo the sample of high activity level populations, males have moe cuved femoa than do females. This diffeence disappeas when the whole sample is consideed and eflects a postulated eduction in division of labou fom the onset of the adoption of agicultue whee both sexes paticipate in agicultual activities (Ruff, 1999). Ruff (1999) suggests the impotance of teain elief on anteoposteio hypetophy of the femoal shaft. Duing downhill walking the impact of the foce is dissipated at incemental angles athe than at a staight angle though the bone esulting in less impact on the joints. The estimation of teain elief fo each of the moden human samples was beyond the scope of this study, but a matix coelation between anteio femoal cuvatue and altitude of the mean location of population (which could 242

274 potentially seve as a poxy fo elief), show that thee is potential to develop this idea futhe. In ode to do this, it would be necessay to include samples in these analyses fo which teain elief and home ange data is available and fo which othe factos such as climate, and activity levels emain constant. These esults suppot the hypothesis that femoal cuvatue is a bone esponse to stesses and stains pesent duing habitual behaviou. opulations with an aquatic subsistence stategy have less biomechanical stess on the lowe limb (Stock and feiffe, 2001; Stock, 2002; Stock, 2006) compaed to the othe subcategoies within high activity goups, and this is shown in that the lowest degee of cuvatue and highest apex of cuvatue in aquatic foageing populations. The pastoalists have the highest teestial mobility and also the highest degee of cuvatue. The esults pesented hee demonstate the potential of femoal cuvatue as a pedicto of activity intensity. Femoal cuvatue may be a bette pedicto than coss-sectional obusticity (Ruff, 1987; eason, 2000b; Ruff and Tinkaus, 2000; Stock, 2002; Stock and feiffe, 2004; Stock, 2006) which is affected also by both activity levels and climate (eason, 2000b; Stock, 2006). Thee wee fou biomechanical hypothese fo long bone cuvatue: 1) cuvatue lowes bending stess by tanslating bending stess to axial compession (Fost, 1967; Hall, 2004), 2) cuvatue facilitates muscle expansion and packing (Lanyon et al., 1979; Lanyon, 1980), 3) cuvatue is a compomise between bone stength and pedictability of bending stains and mateial failue (Lanyon, 1980, 1987; Betam and Biewene, 1988), o 4) geneates stains necessay fo optimal stength (Lanyon, 1980). Out of the fou biomechanical hypotheses fo long bone cuvatue that wee suggested in Chapte 2, it is unlikely that the stess eduction hypothesis (Fost, 1967) accounts fo the diffeences in femoal cuvatue between these human populations as it has been widely demonstated that most of the stess in the long bones is bending stess and that inceased cuvatue is coelated with inceases in bending stess (Lanyon and Baggott, 1976; Lanyon and Boun, 1979; Lanyon et al., 1979; Lanyon, 1980; Biewene, 1983; Lanyon and Rubin, 1986; Lanyon, 1987; Betam and Biewene, 1988; Swatz, 1990; Biewene and Betam, 1994; Main and Biewene, 2004). The second hypothesis suggests that cuvatue facilitates muscle packing (Lanyon, 1980). By inceasing cuvatue the tendons ae able to attach close to the joints while the cuvatue of the 243

275 shaft accommodates the lage bellies of the muscles in the midshaft egion (Lanyon, 1980). Duing ontogeny the development of the muscles on the concave side of the shaft inceases the peiosteal pessue on the shaft which esults in inceased concavity and cuvatue. This hypothesis is suppoted by the adius and tibia of many mammals (Lanyon, 1980), but Swatz (1990) found no coelation between musculatue and cuvatue in anthopoids. Howeve, this study does povide suppot fo the hypothesis in that inceased cuvatue is found in humans fom goups with high activity levels who ae likely to be moe muscula than those fom goups with lowe activity levels The thid hypothesis suggests that a high degee of cuvatue inceases bending moments which ultimately may incease bone stength. Maintaining a modeate amount of stain is necessay fo maintenance of bone mass (Lanyon, 1980; Biewene, 1983; Biewene and Betam, 1994; eason and Liebeman, 2004; Ruff et al., 2006). Theefoe, inceased cuvatue may povide a physiological benefit without affecting second moments of aea o coss-sectional aea (Lanyon, 1980). This is suppoted in the esults fom this study in that high degees of femoal cuvatue tend to be coelated with inceased levels of obusticity. The fouth hypothesis suggests that cuvatue gives pedictability to the diection of bone failue (Lanyon, 1980, 1987; Betam and Biewene, 1988). Because the bone is loaded though bending stess athe than axially when it is cuved, it is pedicted that if a lage amount of stess is applied, the bone is most likely to suffe fom failue (factue) in the diection of the cuvatue. Theefoe, athe than maintaining low amounts of stain by having a staight and axially loaded shaft, cuvatue seves as safety facto of a biological stuctue equiing inceased stength in a single location, athe than acoss the bone (Alexande, 1981). This is suppoted by the esults pesented hee in that individuals with a highe degee of cuvatue have a moe anteoposteioly wide shaft. Results fom studies of coss-sectional obusticity suggest that the cotical bone at midshaft is thickened in the anteoposteio plane (Ruff, 1999), and theefoe in the diection of the cuve, athe than in the mediolateal plane. In ode to fully undestand this inteaction, howeve, it is necessay to combine the cuvatue data with measues of cosssectional geomety. The shape analysis also found that the human femoal shaft shows vaiation in the sinusoidal shape of the lateal side of the shaft. opulations with lowe activity levels had significantly moe sinusoidal femoal shafts compaed to the modeate and high activity goup. These 244

276 diffeences could be due to a decease in bone emodelling ates and lack of physiologically beneficial stains in the shaft. Lowe levels of habitual loading can potentially cause the bone to be less dense and, theefoe, moe susceptible to the pessue of muscles, o cause the bone to take on a sinusoidal shape because thee is no need fo maintaining stuctual integity. The lack of a coelation with othe mophological and behavioual factos makes the sinusoidal aspect of femoal cuvatue difficult to intepet. In summay, the hypotheses discussed above suggest that femoal cuvatue is a esult of inceased activity levels and can be biomechanically explained by facilitating geate muscle mass, geneating physiologically beneficial stains and may incease the pedictability of mateial failue. Lowe am The adius has two cuves which wee used in the analyses. The medial cuve descibes the development of the inteosseous cest, wheeas the lateal cuved descibes the oveall degee of cuvatue of the bone. Thee was no diffeence between populations epesenting the diffeent activity levels fo eithe the medial o the lateal cuve. Howeve, thee wee some diffeences between the subsistence goups. The hoticultualists wee the least cuved. Hoticultualists use thei uppe limbs fo subsistenc-elated activity, though, so this esult cannot be explained by intensity of subsistence-elated activity. It was pedicted that the aquatic foages would have the highest degees of oveall cuvatue. They had a high degee of lateal cuvatue but a low degee of medial cuvatue eflecting the stong development of the inteosseous cest. The aquatic foages also had a poximal medial development on the inteosseous cest and a medially expanded ulna notch. These may eflect the inceased use of the foeam duing the use of watecaft and stonge development of the inteosseous membane. Although none of the shaft shape Cs of the ulna showed diffeences between the activity levels o subsistence goups, the aquatic foages have the longest ulna neck (geatest distance between the tip of the coonoid and the 80% level of the shaft). While it was pedicted that thee would be a coelation between the position of the adial tubeosity and the neck-length of the adius and adial cuvatue these pedictions wee not suppoted by the esults. Thee was no diffeence between males and females and adial cuvatue and ulna shaft shape. 245

277 Although some of the esults suppot the hypothesis that cuvatue is a bone esponse to stesses and stains duing habitual behaviou, the esults ae inconclusive and may be explained by the diffeences in climate instead (see moe below). Hypothesis 3: Cuvatue is a consequence of adaptation to cold climate. Based on Begmann and Allen s ule elated to body size and body popotions, it is known that individuals fom colde climates have shotened distal limbs and that these diffeences ae established though genetic adaptations athe than individual ontogeny (Y'Edynak, 1976; Eveleth and Tanne, 1990; Ruff et al., 1994; eason, 2000b; Van Andel, 2003; Weave, 2003; Ruff et al., 2005). Foeshotening of the limbs may have an effect on cuvatue. Femu As was shown in the past (eason, 2000b; Stock, 2006), thee was a significant coelation between latitude and obusticity. Neithe femoal cuvatue no apex of cuvatue shows any significant pattens with latitude, despite the coelation of latitude with othe mophological featues, such as femoal length and epiphysis size. This would suggest that othe mophological elements that ae unde stong climatic influence, such as the pelvis width, neck-shaft angle (coelated with tosion angle) and body size (fom femoal length) (Ruff, 1995; Weave, 2003) would not be coelated with cuvatue. With the exception of pelvis width, all these vaiables wee exploed, and none wee coelated with eithe degee of cuvatue o position of the apex of cuvatue. Theefoe, femoal cuvatue is not a consequence of adaptation to cold climate. Lowe am Lateal cuvatue of the adius is elated to climate. This is also eflected in the highe degee of lateal cuvatue fo the aquatic foages who have the lowest mean annual tempeatue (Inuit, Russian Eskimo, Geenland Inuit but less Andamanese). The low degee of cuvatue fo the hoticultualists can be explained by the elative wam climate in which these goups live (New Mexico and Ohio). The development of the poximal inteosseous cest of the adius is also highly coelated with climate but is likely a sign of the habitual aquatic subsistence-elated behaviou (such as the use of watecaft). The aquatic foages show a highe adial neck-shaft 246

278 angle which may be elated to the use of the foeam duing the use of watecaft o fishing. Aquatic foages do not stand out in the othe univaiate measuements of the adius o ulna. Robusticity of the distal aticulation of both the ulna and the adius is highly coelated with climate eflecting the elatively shot foeam bones. The adius in populations fom highe latitudes is moe sinusoidal but shows no paticula pattens in the est of its mophology with climate othe than cuvatue. Fo the ulna thee ae some inteesting pattens. Individuals fom highe latitudes have lage poximal ulnas, lage neck-shaft angles (joint-axis angle), a moe infeio insetion of the bachial tubeosity, a smalle distance between 80% of the shaft and the tip of the olecanon, a moe poximoanteio tochlea notch, a moe anteoposteioly sinusoidal shape, a less mediolateal sinusoidal shape and a lage adial notch suface aea. The anteoposteio sinusoidal shape of the ulna is coelated with lateal cuvatue of the adius and eflects the posteio subtense discussed in Fische (1909). An incease in lateal adial cuvatue, a moe sinusoidal adial shaft and the inceased anteoposteio sinusoidal shape of the ulna is likely a consequence of the shotening of the lowe limbs. In the light of the biomechanical hypotheses discussed above fo the femu, cuvatue of the ulna and adius cannot be explained by factos caused by axial loading of the shaft. Cuvatue of the foeam is most likely a way of facilitating muscle packing in esponse to the eduction in elative long bone length in cold-adapted populations. Maintaining the tendon insetions of muscles close to the joints and peventing shotening of the muscles inseting on the shaft (and theefoe pevent loss of contaction function of the ponato tees) aids hoizontal muscle packing by allowing space fo the muscle bellies (Lanyon, 1980). It also maximises the degee of ponation and supination by maintaining the size and axis of otation (Yasutomi et al., 2002). The next step in testing this hypothesis is to combine data fom this study with data on muscle development. The othe biomechanical hypotheses explaining cuvatue of the foeam do not have any diect suppot fom the data in this study. Despite the size of the adial aticula sufaces being coelated with adial cuvatue, thee is no coelation with midshaft obusticity. The coelations between ulna shaft shape and obusticity ae not consistent acoss the bone. Theefoe, these esults ae inconclusive in thei suppot fo the mateial failue pedictability hypothesis (Betam and Biewene, 1988). In ode to test the hypothesis of physiological benefit to the bone, it is necessay to combine the cuvatue data of the foeam sample in this study 247

279 with measues of coss-sectional geomety. Vey little is known about coss-sectional geomety of the ulna and adius and midshaft shape of the adius and extenal shape atios in these moden human samples wee not coelated with lateal o medial cuvatue. 248

280 edicting cuvatue in Neandethals and ealy moden humans. The esults of the vaiation in cuvatue of the femu, ulna and adius within moden humans indicate that thee ae pattens of longitudinal long bone cuvatue but that these ae diffeent fo the uppe and lowe limb. Seveal of the conclusions fom the analyses of ecent humans ae especially elevant and can povide a famewok fo looking at the meaning of long bone cuvatue in alaeolithic samples. Long bone cuvatue follows diffeent tends than obusticity and is not necessaily a esponse to the same types of loading (Ruff et al., 1993; Ruff et al., 1994; Tinkaus et al., 1994; Tinkaus et al., 1999a; eason, 2000b; Ruff and Tinkaus, 2000; Shackelfod and Tinkaus, 2002; Stock, 2002; Stock and feiffe, 2004; Stock, 2006; Calson et al., 2007; Shackelfod, 2007). The highest levels of cuvatue fo the femu wee identified in samples with high activity levels. Theefoe, it is hypothesised that both ealy moden humans and Neandethals will possess high degees of femoal cuvatue and a moe distal apex of cuvatue. The cuvatue of the adius and ulna is stongly influenced by climate. Individuals fom colde climates tend to have moe cuved ulnae and adii. Neandethals, as a goup, wee subject to cold climatic conditions fo a moe extended peiod of time than any moden human population, so it is hypothesised that they have hypepola adaptations to the climate in which they lived (Boule and Vallois, 1952; Tinkaus, 1981; Chuchill, 1998; eason, 2000a, 2000b; Aiello and Wheele, 2003; Weave, 2003; Kause et al., 2007; Shackelfod, 2007). Hence, the Neandethals adius is pedicted to have a highe degee of lateal cuvatue and a moe sinusoidal shape, and the ulna is pedicted to be moe anteoposteioly sinusoidal than any othe moden human sample. The ealy moden humans ae pedicted to have less adial and ulna cuvatue than Neandethals as they wee not exposed to the cold Euopean climate fo the same extended time. Depending on the time the ealy moden humans spent in the cold Euopean climate, it can be hypothesised that they, too, may have a high degee of cuvatue. Howeve, as moden humans wee likely to have oiginated in topical Afica (Mellas and Stinge, 1989; Smith et al., 1989; Ba-Yosef, 1992; Deacon, 1992; Stinge, 1992; Ingman et al., 2000; eason, 2000a; Stinge, 2002; White et al., 2003; Mellas, 2004) they may display vey low levels of cuvatue. These hypotheses will be tested in Chapte

281 CHATER 5. LONG BONE CURVATURE IN NEANDERTHALS, EARLY AND RECENT MODERN HUMANS Objective The pupose of the intespecific analyses is to detemine whee fossil specimens fall elative to pattens of vaiation in long bone cuvatue in ecent moden humans. The Neandethal and ealy moden human fossil specimens ae included in the Geneal ocustes Analyses and in the incipal Component analyses. The inclusion of the fossils in the incipal Component analysis slightly changes the distibution of the shape changes along the pincipal components and will be discussed below. In ode to examine vaiation in long bone mophology, the pincipal component scoes ae used in Analyses of Vaiance (ANOVAs) and post-hoc tests using paiwise compaisons. As in Chapte 4, the Hochbeg GT2 and Games-Howell pocedues wee used (discussed in moe detail in Chapte 3: Mateials and Methods) and the esults will be discussed fo significant Fscoes. Fo these analyses, fossil hominins ae eithe gouped as Neandethals o ealy moden humans. To detemine the elationship between the diffeent aspects of mophology and goup diffeences, disciminant functions ae calculated using the pincipal component scoes as independent vaiables. In the esults descibed below the abbeviations of the pincipal components (Cs) names ae made up of thee pats. The fist designates the landmak set included in the study (i.e. acuve stands fo anteio cuve). The second designates the sample included (i.e. ALL stands fo all fossils and all ecent moden humans). The thid is the C numbe (i.e. C2 stands fo second C), e.g. AcuveAllC1. 250

282 5.2. Femu Femu pincipal components explained As was the case fo the investigation of intaspecific vaiation within moden humans, the changes fo each of the cuves and the poximal and distal epiphyses (epiphyses) along each pincipal component ae visualised using Mophologika. Although the Cs ae vey simila to those obtained when only the moden humans ae consideed, thee ae diffeences between the C scoes and shape changes along the Cs. Theefoe, the Cs will be explained again below. The figues pesented coespond to the most exteme positive and negative individuals on the scale fo each C. The cuves ae semi-landmaks only, wheeas the epiphyses ae landmaks. Viewing angles wee chosen to illustate similaities and diffeences most clealy. Fo the cuves, this is in lateal view, unless othewise stated in the Figue captions. The Neandethal sample consists of eight specimens, the ealy moden humans sample consists of 13 specimens, and 428 individuals ae included in the ecent moden human sample Anteio suface (acuve) The fist thee pincipal components explain 63.7%, 9.62%, and 7.30% of the vaiance,,espectively,(total 80.06%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. AcuveAllC1 eflects vaiation in degee of anteio cuvatue o subtense (Figue 5-1a). The second pincipal component (acuveallc2) is elated to the position of the apex of cuvatue (Figue 5-1b). The thid pincipal component is the shape of the shaft in anteio view (Figue 5-1c). Negative values ae moe sinusoidal, wheeas positive values ae staight. 251

283 a b c Figue 5-1 Mophological tends fo the anteio cuve of the femu fo Neandethals, ealy and ecent moden humans. a: incipal component 1: lateal view. Negative values ae less cuved, positive values ae moe cuved. b: incipal component 2: lateal view. Individuals with negative values have a moe poximal apex of cuvatue, wheeas those with positive values have a moe distal apex of cuvatue. c: incipal component 3: anteio view. Negative values ae the staightest, wheeas positive values indicate a mediolateally sinusoidal shape. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 252

284 osteio suface (pcuve) The fist fou pincipal components of the posteio cuve analysis explain 34.9%, 14.8%, 11.4% and 7.47%,,espectively, of the vaiation (total 68.5%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. The posteio cuve is vey simila to the anteio cuve. cuveallc1 eflects diffeences in degee of cuvatue o subtense (Figue 5-2a) (note that pcuveallc1 is loaded in an opposite diection fom the othe cuvatue Cs). The second pincipal component (pcuveallc2) is the shape of the shape of the cuve in posteio view (Figue 5-2b). The thid pincipal component (pcuveallc3) is elated to the apex of the posteio cuve (Figue 5-2c). The fouth pincipal component (pcuveamhc4) is the diection of the distal end of the cuve (Figue 5-2d). 253

285 a b c d Figue 5-2 Mophological tends fo the posteio cuve of the femu fo Neandethals, ealy and ecent moden humans. a: incipal component 1: lateal view. Negative values ae moe cuved, positive values ae less cuved. b: incipal component 2: anteio view. Negative values ae the staightest, wheeas positive values ae mediolateally sinusoidal. c: incipal component 3: lateal view Negative values have a highe apex of cuvatue compae to positive values. d: incipal component 4: lateal view. ositive individuals have a moe posteioly pojected distal cuve. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 254

286 Medial suface (mcuve) The fist thee pincipal components of the medial cuve analysis explain 49.9%, 16.6%, and 15.39%,espectively,of the vaiation (total 83.1%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. As was the case in the analysis on moden humans the component mcuveallc1 eflects diffeences in degee of anteio cuvatue (Figue 5-3a). The second pincipal component (mcuveallc2) is elated to the position of the apex of cuvatue (Figue 5-3b). The thid pincipal component (mcuveallc3) is the posteio extension of the distal end of the cuve and the evenness of the cuve (Figue 5-3c). 255

287 a b c Figue 5-3 Mophological tends fo the medial cuve of the femu fo Neandethals, ealy and ecent moden humans. All lateal view. a: incipal component 1. Negative values ae less cuved, positive values ae moe cuved. b: incipal component 2. Negative values have a highe apex of cuvatue compae to positive values. c: incipal component 3. ositive values ae moe flattened off with inceased posteio pojection of the distal cuve, wheeas negative values have a cuve that appoximates an ac of a cicle. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 256

288 Lateal suface (lcuve) The fist fou pincipal components of the lateal cuve analysis explain 51.3%, 15.5%, 9.54% and 5.44%,,espectively, of the vaiation (total 81.78%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. The component lcuveallc1 eflects diffeences in anteio cuvatue o subtense (Figue 5-4a). The second pincipal component (lcuveallc2) is elated to the position of the apex of cuvatue and the diection of the poximal pat of the suface (Figue 5-4b). The thid pincipal (lcuveallc3) component is elated to the staightening of the femu at the level of the lesse tochante (Figue 5-4c). The fouth pincipal component (lcuveallc4) is the shape of the lateal suface in anteio view (Figue 5-4d). 257

289 a b c d Figue 5-4 Mophological tends fo the lateal cuve of the femu fo Neandethals, ealy and ecent moden humans. a: incipal component 1: lateal view. Negative values ae less cuved, positive values ae moe cuved. b: incipal component 2: lateal view. Negative values have a moe distal apex of cuvatue and little posteio diection of the poximal cuve, wheeas those with positive values have a moe poximal apex of cuvatue and a moe posteioly pojecting poximal cuve. c: incipal component 3: lateal view. ositive values show a flattening off at the level of the lesse tochante and negative values ae evenly cuved. d: incipal component 4: anteio view. ositive values ae the staightest, wheeas negative values have a mediolateally sinusoidal shape. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 258

290 oximal and distal epiphyses (Epi) The fist five pincipal components of the epiphysis analysis explain 14.5%, 9.62%, 7.47%, 5.30% and 4.34%,,espectively, of the vaiation (total 43.9%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. The component epiallc1 eflects diffeences in the width of the distal epiphyses and the neckshaft angle (Figue 5-5a). The second pincipal component (epiallc2) is elated to the oveall width of the femu and the position of the lesse tochante (Figue 5-5b). The thid pincipal component (epiallc3) is elated to the width of the distal epiphyses and degee of tosion (Figue 5-5c). The fouth pincipal component (epiallc4) is had to intepet and it is unclea what it elates to. The changes along the pincipal component axis ae vey subtle and this C will not be included in any of the the subsequent analyses. The fifth pincipal component (epiallc5) elates to the length of the neck and the depth of the distal epiphyses (Figue 5-5d). 259

291 a b c d 260

292 Figue 5-5 Mophological tends fo the epiphyses of the femu fo Neandethals, ealy and ecent moden humans. a: incipal component 1: anteio view. Individuals with negative values have wide distal epiphyses and a lowe neck-shaft angle. b: incipal component 2: anteio view. Negative values have wide distal epiphyses and heads and a lowe lesse tochante, wheeas positive values ae naow and have a elatively highe lesse tochante. c: incipal component 3: supeio view. Individuals with negative values have wide distal epiphyses and less tosion than those with positive values. d: incipal component 5. Individuals with negative values have a long neck and deep knees compaed to individuals with positive values. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C Summay As in the analysis of ecent moden human populations, anteio cuvatue is the most impotant pincipal component in all fou cuves (acuveallc1, pcuveallc1, mcuveallc1, lcuveallc1). This is eflected in the significant coelations between all these cuves (note that pcuveallc1 is loaded in the opposite diection fom the othe cuvatue Cs and is theefoe negatively coelated with them) (Table 5-1). Fo this eason, only acuveallc1 and pcuveallc1 will be analysed and discussed. The position of the apex of cuvatue is the majo facto in acuveallc2, pcuveallc3, mcuveallc2 and lcuveallc2, so only acuveallc2 and pcuveallc3 will be discussed. These ae also all coelated, but none of the -values ae high (Table 5-2). The othe pincipal components fo the cuves explain mino changes in suface shape and will be included in subsequent analyses. 261

293 Table 5-1 eason s coelation matix: femoal cuvatue Cs (n= 449). Neandethals, ealy and ecent moden humans. AcuveAllC1 pcuveallc1 McuveAllC1 pcuveallc ** <0.001 McuveAllC ** ** <0.001 <0.001 LcuveAllC ** ** 0.368** <0.001 <0.001 <0.001 * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). Table 5-2 eason s coelation matix: femoal apex of cuvatue Cs (n= 449). Neandethals, ealy and ecent moden humans. AcuveAllC2 pcuveallc3 McuveAllC ** <0.001 McuveAllC ** 0.152** <0.001 <0.001 LcuveAllC ** 0.177** 0.213** <0.001 <0.001 <0.001 * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). pcuveallc3 262

294 Diffeences in femoal mophology between Neandethals, ealy and ecent moden humans Cuvatue The goups ae significantly diffeent fo both cuvatue Cs: acuveallc1, pcuveallc1 (Table 5-3). Neandethals have the highest degee of anteio and posteio cuvatue, followed by ealy moden humans. Recent moden humans ae the staightest (Figue 5-6 and Figue 5-7). Statistically, Neandethals ae diffeent fo both pincipal components fom ecent moden humans. Ealy moden humans ae diffeent fom Neandethals fo acuvallc1 only (Appendix 39). Box plots ae used in ode to display cuvatue and apex of cuvatue fo the sepaate fossils. Table 5-3 ANOVA esults fo palaeogoup1 and femoal cuvatue Cs. d.f.=2 AcuveAllC1 pcuveallc1 *=significant at α=0.05 F Sig. <0.001* <0.001* 1 alaeogoup efes to the thee categoies commonly used in palaeoanthopological eseach that ae included in these analyses: Neandethals, ealy anatomically moden humans, ecent moden humans. 263

295 Figue 5-6 The anteio cuve of the femu fo Neandethals, ealy and ecent moden humans. (Line=mean, Box= 2 S.E., whiskes: 2 S.D.). The highe values fo Neandethalss indicate that they ae moe cuved than the moden humans. Figue 5-7 The posteio cuve of the femu fo Neandethals, ealy and ecent moden humans. (Line=mean, Box= 2 S.E., whiskes: 2 S.D.). The lowe values fo Neandethal indicates that they ae moe cuved than the moden humans. 264

296 Apex of cuvatue The goups ae significantly diffeent fo the position of the apex of cuvatue in one C (Table 5-4). On the anteio suface, Neandethals have the lowest apex of cuvatue and ae significantly diffeent fom ealy and ecent moden humans (Figue 5-8 and Appendix 40). Table 5-4 ANOVA esults fo palaeogoup and femoal apex of cuvatue Cs. d.f.=2 AcuveAllC2 pcuveallc3 *=significant at α=0.05 F Sig * Figue 5-8 The anteio apex of cuvatue of the femu fo Neandethals, ealy and ecent moden humans. (Line=mean, Box= 2 S.E., whiskes: 2 S.D.). The highe value fo Neandethals indicates a lowe apex of cuvatue. 265

297 Othe shaft shape The goups ae significantly diffeent fo only one of the othe shaft shape Cs (Table 5-5). The post-hoc tests shows that Neandethals may have a lateal cuve that staightens at the level of the lesse tochante (lcuveallc3)(appendix 41). Table 5-5 ANOVA esults fo palaeogoup and othe femoal shaft shape Cs. d.f.=2 AcuveAllC3 pcuveallc2 pcuveallc4 McuveAllC3 LcuveAllC3 LcuveAllC4 *=significant at α=0.05 F Sig * Figue 5-9 LcuAllC3 fo Neandethals, ealy and ecent moden humans. The lowe values fo Neandethals indicate wide distal epiphyses and a lowe neck-shaft angle. Mean and 95% confidence inteval (whiskes). 266

298 Epiphysis shape Neandethals have moe obust epiphyses, a lowe neck-shaft angle and a lowe lesse tochante than ealy and ecent moden humans (EpiAllC1 and EpiAllC2) (Table 5-6 and Figue 5-10 and Figue 5-11; Appendix 42). Neandethals also have less tosion (EpiAllC3) (Figue 5-12) and a long neck and deep distal condyles (EpiAllC5) (Figue 5-13). Table 5-6 ANOVA esults fo palaeogoup and othe femoal shaft shape Cs. d.f.=2 F EpiAllC EpiAllC EpiAllC EpiAllC *=significant at α=0.05 Sig. <0.001* 0.003* 0.043* 0.008* Figue 5-10 EpiAllC1 fo Neandethals, ealy and ecent moden humans. The lowe values fo Neandethals indicate wide distal epiphyses and a lowe neck-shaft angle. Mean and 95% confidence inteval (whiskes). 267

299 Figue 5-11 EpiAllC2 fo Neandethals, ealy and ecent moden humans. The lowe values fo the Neandethals indicate wide distal epiphyses and heads and a lowe lesse tochante. Mean and 95% confidence inteval (whiskes). Figue 5-12 EpiAllC3 fo Neandethals, ealy and ecent moden humans. The lowe values fo Neandethals indicate wide distal epiphyses and less tosion than moden human goups with positive values. Mean and 95% confidence inteval (whiskes). 268

300 Figue 5-13 EpiAllC5 fo Neandethals, ealy and ecent moden humans. The lowe values fo Neandethals indicate a long neck and anteoposteioly deep distal epiphyses. Mean and 95% confidence inteval (whiskes) Univaiate measuements The goups ae significantly diffeent fo all univaiate measuements (Figue 5.7). The highest F-scoes ae fo head-obusticity, neck-length, neck-shaft angle and obusticity index (Table 5-8). ost-hoc tests (Appendix 43) indicate that Neandethals have the lagest femoal head, longest neck and lagest distal epiphyses compaed to ealy and ecent moden humans, although thei midshaft obusticity and neck-shaft angle is compaable to that of ealy moden humans (Figue 5-23; Figue 5-21 and Figue 5-15). Ealy moden human femoa ae longe and have lowe tosion angles, ae moe obust, and have highe midshaft and subpilastic atios than do ecent moden humans (Figue 5-14; Figue 5-16; Figue 5-22; Figue 5-18; Figue 5-19). Ealy moden human femoa have a high midshaft shape atio, which pobably eflects the stong expession of the linea aspea. Neandethal femoa have an almost ound shaft at the midshaft level and lack a clea linea aspea (Figue 5-18). 269

301 Table 5-7 Desciptives fo Neandethals, ealy and ecent moden humans and the univaiate measuements of the femu. Femu length Neck-shaft angle Tosion angle subtochatio midshaftatio subpilatio condylediamatio necklengthatio obustindex headob Neandethal Ealy Homo sapiens Recent Homo sapiens Neandethal Ealy Homo sapiens Recent Homo sapiens Neandethal Ealy Homo sapiens Recent Homo sapiens Neandethal Ealy Homo sapiens Recent Homo sapiens Neandethal Ealy Homo sapiens Recent Homo sapiens Neandethal Ealy Homo sapiens Recent Homo sapiens Neandethal Ealy Homo sapiens Recent Homo sapiens Neandethal Ealy Homo sapiens Recent Homo sapiens Neandethal Ealy Homo sapiens Recent Homo sapiens Neandethal Ealy Homo sapiens Recent Homo sapiens N Mean S.D Table 5-8 ANOVA esults fo palaeogoup and femoal univaiate measuements. d.f.=2 Femu length Neck-shaft angle Tosion angle subtochatio midshaftatio subpilatio condylediamatio necklengthatio obustindex headob *=significant at α=0.05 F Sig * <0.001* 0.001* 0.005* 0.008* 0.007* 0.001* <0.001* <0.001* <0.001* 270

302 Figue 5-14 Femu length fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). Figue 5-15 Neck-shaft angle fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 271

303 Figue 5-16 Tosion angle fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). Figue 5-17 Subtochanteic atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 272

304 Figue 5-18 Midshaft atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). Figue 5-19 Subpilastic atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 273

305 Figue 5-20 Condyle obusticity fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). Figue 5-21 Neck length atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 274

306 Figue 5-22 Robusticity index fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). Figue 5-23 Head obusticity fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 275

307 Disciminant function analysis A DFA with coss-validation using all Cs (included in the above analyses) and univaiate measuements used in the analyses above was used to sepaate Neandethals, ealy and ecent moden humans. Function 1 sepaates best between Neandethals and moden humans in geneal, wheeas function 2 sepaates ealy moden humans fom ecent moden humans (Figue 5-24). The vaiables in Table 5-9 appea in the ode of thei disciminating powe. Function 1 eflects (odeed accoding to deceasing coelation between the vaiable and the function) degee of cuvatue, obusticity of the head, width of the distal and poximal femu, neck-length atio, low neck-shaft angle, obusticity. Function 2 eflects the midshaft and subpilastic shaft shape, femu length, and othe aspects of shaft shape (Table 5-9). Fo these thee populations (Neandethals, ealy and ecent moden humans) with vey uneven sample sizes, the expected popotion of coect andom classification based on sample size is ~90%. The DFA with coss-validation was able to coectly classify Neandethals and ecent moden humans elatively succesfully with 87.5% (7 out of 8 Neandethals) and 99.5% (425 out of 427 moden humans) classified coectly. Ealy moden humans wee almost all classified as ecent moden humans (92.3% - 12 out of 13), although none wee classified as Neandethals. Oveall, fo the thee goups togethe, this gives 96.7% of coect classification. 276

308 Function 2 Function 1 Figue 5-24 Disciminant Function 1 and 2 fo Neandethals, ealy and ecent moden humans. 277

309 Table 5-9 Disciminant function coefficients - femu. Function 1 pcuveallc1 AcuveAllC1 headob EpiAllC1 necklengthatio Neck-shaft angle LcuveAllC1 obustindex condylediamatio EpiAllC2 Tosion angle subtochatio EpiAllC3 McuveAllC1 AcuveAllC3 Canonical R= Λ = <0.001 Function 2 midshaftatio subpilatio Femu length AcuveAllC2 EpiAllC4 LcuveAllC2 LcuveAllC4 pcuveallc4 McuveAllC3 EpiAllC5 pcuveallc2 AcuveAllC4 McuveAllC2 pcuveallc3 Canonical R= Λ = <

310 Summay Neandethals have femoa with a highe degee of anteio cuvatue than do ealy moden humans and ecent moden humans. They also have the most distal apex of cuvatue. They have wide and deepe distal epiphyses, lage femoal heads, lowe neck-shaft angles (only compaed to ealy moden humans) and ae the most obust (significantly diffeent fom ecent moden humans only). Disciminant function classification vey successfully distinguished Neandethals fom the ecent moden human goups, but the ovelap between ealy and ecent moden humans esulted in fequent misclassification of ealy moden humans into the much lage and moe vaiable ecent moden human goup. 279

311 5.3. The lowe am The esults pesented hee will fist discuss the pincipal components and visualisations, using the same appoach that was used fo the section on the femu. The adius sample consists of 15 Neandethals, 15 ealy moden humans and 361 ecent moden humans. The ulna sample consists of 13 Neandethals, 21 ealy moden humans and 344 ecent moden humans Radius shape pincipal components explained Medial suface (mcuve) The fist thee Cs of the medial cuve explain 46.1%, 13.2% and 8.94%, espectively, of the vaiation (total 68.2%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. C1 eflects the vaiation in lateal cuvatue of the adius (Figue 5-25a). C2 is elated to the medial expansion of the poximal inteosseous cest and the diection of the distal end of the medial suface (Figue 5-25b). C3 is the sinusoidal shape of the shaft in the anteoposteio plane (Figue 5-25c). 280

312 a b c Figue 5-25 Mophological tends fo the medial cuve of the adius fo Neandethals, ealy and ecent moden humans. a: incipal component 1: anteio view. Negative values have a highe degee of cuvatue than positive values. b: incipal component 2: anteio view. ositive values show an inceased medial extension of the poximal inteosseous cest and a medial diection of the distal cuve (moe medially expanded ulna notch), wheeas negative values show no medial expansion of the poximal inteosseous cest and an ulna notch that is not medially pojected. c: incipal component 3: lateal view. ositive values have a moe sinusoidal shape, wheeas negative values have no sinusoidal shape. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 281

313 Lateal cuve (lcuve) The fist thee Cs of the lateal cuve explain 40.6%, 20.9% and 9.43%,espectively,of the vaiation (total 70.9%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. Simila to the analyses on moden humans C1 eflects diffeences in lateal cuvatue (Figue 5-26a). C2 is influenced by the apex of cuvatue and the diection of the distal end of the lateal suface (Figue 5-26b). C3 elates to the sinusoidal shape of the lateal cuve in the anteoposteio plane (Figue 5-26c). 282

314 a b c Figue 5-26 Mophological tends fo the lateal cuve of the adius fo Neandethals, ealy and ecent moden humans. a: incipal component 1: anteio view. Negative values have a highe degee of cuvatue wheeas positive values have a lowe degee of lateal cuvatue. b: incipal component 2: anteio view. ositive values have a moe poximal apex of cuvatue and a moe lateally pojecting styloid pocess, wheeas negative values have thei apex of cuvatue at midshaft and lack the lateal pojection of the styloid pocess. c: incipal component 3: lateal view. ositive values ae moe sinusoidal. Negative values ae not sinusoidal. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 283

315 Epiphyses (Epi) The fist two C s of the epiphysis analysis explain 33.3% and 8.53%,,espectively, of the vaiation (total 41.8%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. When scatteplots of the Cs wee obseved, C6 (4.71% of vaiation) showed Neandethals to have pimaily positive values and is theefoe included in the following analyses. C1 eflects the diection of the head and the distal aticula suface in elation to the shaft (Figue 5-27a). C2 elates to the length of the adius between the adial tubeosity and 80% level of the shaft and the oientation of the tip of the styloid pocess (Figue 5-27b). C6 is elated to the position of the adial tubeosity (Figue 5-27c). 284

316 a b c Figue 5-27 Mophological tends fo the epiphyses of the adius fo Neandethals, ealy and ecent moden humans. All medial view. a: incipal component 1. Individuals with negative values have a moe anteioly oiented head, wheeas those with positive values ae moe posteioly oiented. b: incipal component

317 Negative values indicate a shote distance between the adial tubecle and the 80% level of the shaft and a moe posteioly located styloid pocess and positive values have a longe distance and a moe anteioly located styloid pocess. c: incipal component 6. Individuals with negative values have a moe anteioly located adial tubeosity compaed to those with positive values who have a moe posteioly located tubeosity. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C Summay Degee of mediolateal cuvatue is the most impotant C fo both medial and lateal suface (mcuveallc1 and lcuveallc1). This is eflected in the significant coelation (=0.369) between the two cuvatue Cs. Thee is no significant coelation between the Cs of the epiphyses and the two cuvatue Cs (Table 5-10 and Table 5-11). Coelations between the othe shaft shape Cs indicate that individuals who have a lowe degee of medial cuvatue (mcuveallc1) have an apex of cuvatue at midshaft and a weakly developed styloid pocess (lcuveallc2), and a less sinusoidal shaft (lcuveallc3) (Table 5-12). A highe degee of lateal cuvatue is elated to an inceased development of the poximal inteosseous cest and inceased medial pojection of the adial notch (mcuvec2). Thee is no coelation between the epiphysis and the othe shaft shape Cs (Table 5-13). 286

318 engevnah shanida1 dolni pavlovt combcplt Ohio kebaalt shanida6 sungit egout LcuAllC neandet feas1lt cfeaslt cfeast Tasmanian dolni16l Hottentot South Dakota Andaman New Mexico Bantou Austalian Coloado native oint Hope Alaska patau2lt Tiea del Fuego combcpt Natufian feas2t Russian Mesolithic Sibeia ohalo21 ygmee dolni13l Alaskan Aleut kostie14 Kazach Geenland Inuit Russian Eskimo kebaat Lapland dolni14l chapelle sgemain Qafzeh09 skhul4lt feas1t spynean1 moustie quina5lt McuAllC1 Figue 5-28 The fist Cs fo the medial and lateal cuve of the adius. Both Cs eflect the degee of mediolateal cuvatue (negative values ae moe cuved). All Neandethals, ealy moden humans and ecent moden human samples. Table 5-10 eason s coelation matix fo adius cuvatue Cs (n= 391). McuAllC1 LcuAllC <0.001** * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). 287

319 Table 5-11 eason s coelation matix fo adius cuvatue and epiphyses Cs (n= 377). EpiAllC1 ApiAllC LcuAllC * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). McuAllC1 Table 5-12 eason s coelation matix fo adius cuvatue and othe shaft shape Cs (n= 391). McuAllC1 LcuAllC1 McuAllC ** McuAllC LcuAllC ** < LcuAllC ** < * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). Table 5-13 eason s coelation matix fo adius epiphyses and othe shaft shape Cs (n= 377). EpiAllC1 EpiAllC McuAllC LcuAllC LcuAllC * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). McuAllC2 288

320 The ulna pincipal components explained osteio cuve (pcuve) The fist fou Cs of the posteio cuve analysis explain 33.7%, 23.3%, 13.4% and 6.31%,,espectively, of the vaiation (total 76.71%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. C1 eflects diffeences in mediolateal cuvatue (Figue 5-29a). C2 is the sinusoidal shape of the shaft in the mediolateal plane (Figue 5-29b). C3 elates to the sinusoidal shape of the lateal cuve in the anteoposteio plane (Figue 5-29c). C4 is the deflection of the poximal shaft (Figue 5-29d). 289

321 a b c d Figue 5-29 Mophological tends fo the posteio cuve of the ulna fo Neandethals, ealy and ecent moden humans. a: incipal component 1: anteio view. Negative values have a highe degee of mediolateal cuvatue, wheeas positive values have a lowe degee of cuvatue. b: incipal component 2: anteio view. ositive values have less of a sinusoidal shape in the mediolateal plane than negative values. c: incipal component 3: medial view. ositive values ae moe sinusoidal compaed to negative values. d: incipal component 4: medial view. ositive values show a bent poximal shaft indicating a moe anteioly pojected head, wheeas negative values ae elatively staight. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 290

322 oximal ulna (pox) The fist thee Cs of the lateal cuve analysis explain 20.4%, 16.6% and 7.89%,,espectively,,of the vaiation (total 44.9%). Subsequent Cs explain minimal amounts of the vaiation and ae not consideed futhe. C1 eflects diffeences in the oientation of the poximal ulna in elation of the shaft (Figue 5-30a). C2 elates to the distance between the 80% level of the shaft and the coonoid pocess (Figue 5-30b). C3 shows the oientation of the tochlea notch (Figue 5-30c). 291

323 a b c Figue 5-30 Mophological tends fo the poximal ulna fo Neandethals, ealy and ecent moden humans. a: incipal component 1: anteio view. ositive values have a poximal ulna that is medially pojected with a medial facing tochlea notch, wheeas negative values have a poximal ulna that is lateally pojected and has a moe lateal facing tochlea notch. b: incipal component 2: anteio view. ositive values have a longe distance between the 80% and the coonoid pocess, wheeas negative values have shot distances. c: incipal component 3: lateal view. ositive values have a moe poximo-anteio facing tochlea notch and negative values have a moe anteio facing tochlea notch. ositive and negative visualisations coespond to the most exteme positive and negative scoes fo each C. 292

324 Summay Thee is no significant coelation between the shaft Cs no ae the poximal ulna Cs significantly elated (Table 5-14). The coelations between the posteio cuve and the poximal ulna Cs showed that individuals with a geate distance between the 80% level of the shaft and the coonoid pocess (poxallc2) have a moe sinusoidal shaft shape in the anteoposteio plane (pcuveallc3). Also, individuals with a moe poximo-anteio tochlea notch (poxallc3) have a less mediolateally sinusoidal shaft shape (pcuveallc2). Table 5-14 eason s coelation matix: ulna Cs (n= 344). pcuallc1 pcuallc2 pcuallc3 pcuallc pcuallc pcuallc oxallc * oxallc ** < oxallc ** < * = Coelation is significant at the 0.05 level (2-tailed). ** = Coelation is significant at the 0.01 level (2-tailed). pcuallc4 oxallc1 oxallc

325 Diffeences in lowe am mophology between Neandethals, ealy and ecent moden humans Cuvatue The goups ae significantly diffeent fo both cuvatue Cs: mcuveallc1 and lcuveallc1 (Table 5-15). Neandethals have a highe degee of lateal and medial cuvatue than ealy and ecent moden humans (Figue 5-31 and Figue 5-32). The ealy and ecent moden human samples ae not diffeent fom each othe (Appendix 44). Table 5-15 ANOVA esults fo palaeogoup and adius cuvatue Cs. d.f.=2 McuAllC1 LcuAllC1 *=significant at α=0.05 F Sig. <0.001* 0.003* Figue 5-31 The medial cuve of the adius fo Neandethals, ealy and ecent moden humans. (Line=mean, Box= 2 S.E., whiskes: 2 S.D.). The lowe values fo Neandethal adii indicate that they ae moe cuved than those of moden humans. 294

326 Figue 5-32 The lateal cuve of the adius fo Neandethals, ealy and ecent moden humans t. (Line=mean, Box= 2 S.E., whiskes: 2 S.D.). The lowe values fo Neandethal adii indicate that they ae moe cuved than those of moden humans Othe shaft shape Fo the adius, the goups ae significantly diffeent fo the lateal shaft shape Cs only (Table 5-16). Neandethals have an apex of cuvatue at midshaft and lack a lateal pojection of the styloid pocess compaed to those of moden humans, who have a moe poximal apex and a moe pojecting styloid pocess (lcuveallc2) (Figue 5-33). Neandethals also have a moe sinusoidal adius in the anteoposteio plane compaed to that of moden humans (lcuveallc3) (Figue 5-34) (ealy moden humans only significantly diffeent using Hochbeg s GT2 (Appendix 44). 295

327 Table 5-16 ANOVA esults fo palaeogoup and othe adius shaft shape Cs. d.f.=2 McuAllC2 McuAllC3 LcuAllC2 LcuAllC3 *=significant at α=0.05 F Sig <0.001* <0.001* Figue 5-33 LcuAllC2 fo Neandethals, ealy and ecent moden humans. The highe values fo the Neandethals indicate an apex of cuvatue at midshaft and a lack of lateal pojection of the styloid pocess. Mean and 95% confidence inteval (whiskes). 296

328 Figue 5-34 LcuAllC3 fo Neandethals, ealy and ecent moden humans. The highe values fo the Neandethals indicate a moe sinusoidal adius in the anteoposteio plane. Mean and 95% confidence inteval (whiskes). Fo the ulna, the goups ae significantly diffeent fo two of the shaft shape Cs (Table 5-17). Neandethals have less mediolateal cuvatue of the ulna shaft compaed to ealy and ecent moden humans (pcuveallc1) (Figue 5-35). Neandethals also have a less mediolateal sinusoidal ulna shaft shape compaed to ecent moden humans (Figue 5-36) (Appendix 46). Table 5-17 ANOVA esults fo palaeogoup and ulna shaft shape Cs. d.f.=2 pcuveallc1 pcuveallc2 pcuveallc3 pcuveallc4 *=significant at α=0.05 F Sig * <0.001*

329 Figue 5-35 cuallc1 fo Neandethals, ealy and ecent moden humans. The highe values fo the Neandethals indicate less mediolateal cuvatue of the ulna shaft. Mean and 95% confidence inteval (whiskes). Figue 5-36 cuallc2 fo Neandethals, ealy and ecent moden humans. The highe values fo the Neandethals indicate a less mediolateal sinusoidal ulna shaft shape. Mean and 95% confidence inteval (whiskes). 298

330 Epiphyses shape The goups ae simila in thei adial epiphyseal shape (Table 5-18). Fo the ulna, the goups ae significantly diffeent fo two poximal shape Cs (Table 5-19). Neandethals have a shote distance between the 80% level of the shaft and the coonoid pocess compaed to ealy moden humans (oxallc2) (Figue 5-37). Thee is a tend fom Neandethals to ecent moden humans in having a moe poximo-anteio athe than an anteio facing tochlea notch and all goups ae significantly diffeent fom each othe (oxallc3) (Appendix 47) (Figue 5-38). Table 5-18 ANOVA esults fo palaeogoup and adius epiphysis shape Cs. d.f.=2 EpiAllC1 EpiAllC2 EpiAllC6 *=significant at α=0.05 F Sig Table 5-19 ANOVA esults fo palaeogoup and poximal ulna Cs. d.f.=2 oxallc1 oxallc2 oxallc3 *=significant at α=0.05 F Sig * <0.001* 299

331 Figue 5-37 oxallc2 fo Neandethals, ealy and ecent moden humans. The lowe values fo the Neandethals indicate a shote distance between the 80% level of the shaft and the coonoid pocess. Mean and 95% confidence inteval (whiskes). Figue 5-38 oxallc3 fo Neandethals, ealy and ecent moden humans. The lowe values fo the Neandethals indicate a moe poximo-anteio athe than an anteio facing tochlea notch. Mean and 95% confidence inteval (whiskes). 300

332 Univaiate measuements The goups ae significantly diffeent fo most univaiate measuements of both the adius and the ulna (Table 5-20 and Table 5-21). Recent moden humans have shote adii than ealy moden humans (Figue 5-39). Neandethals have a moe mediolateally located adial tubeosity than ealy and ecent moden humans (Figue 5-40). Neandethals have a highe degee of dosal and lateal subtense, a longe adial neck and a moe anteoposteioly wide adial head than ealy and ecent moden humans (Figue 5-41; Figue 5-42; Figue 5-43). The ealy moden humans ae simila to the ecent moden humans fo those featues. The midshaft shape atio shows no diffeence between the samples, but a downwad tend in the means suggests a tend towad moe anteoposteio flattening and mediolateal widening which can be intepeted as the inceased development of the inteosseous cest with time (Figue 5-44) (Appendix 48). Table 5-20 Desciptives of palaeogoup and the univaiate measuements of the adius. Max_ Length neck-shaft angle osradtubml DosalST LatealST NeckLengthRatio HeadShapeRatio midshaftshaperation Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human N Mean S.D

333 Table 5-21 ANOVA esults fo palaeogoup and univaiate measuements of the adius. d.f.=2 Max_ Length neck-shaft angle osradtubml DosalST LatealST NeckLengthRatio HeadShapeRatio midshaftshaperatio *=significant at α=0.05 F Sig * * <0.001* <0.001* 0.006* <0.001* 0.023* Figue 5-39 Maximum adius length fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 302

334 Figue 5-40 osition of the adial tubeosity fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). Figue 5-41 Dosal subtense fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 303

335 Figue 5-42 Relative adius neck length fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). Figue 5-43 Head shape atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 304

336 Figue 5-44 Midshaft shape atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). Fo the ulna, thee ae diffeences among the goups fo most univaiate measuements (Table 5-22 and Table 5-23). Neandethals have a elatively lage poximal ulna, smalle shaftolecanon angle, moe even coonoid-olecanon atio and a low bachial tubeosity compaed to both ealy and ecent moden humans (Figue 5-46; Figue 5-49; Figue 5-50; Figue 5-51; Figue 5-52). Neandethals ae also moe obust at midshaft, have a small adial notch, and have a moe anteoposteioly wide ulna shaft than do ecent moden humans (Figue 5-53; Figue 5-48; Figue 5-47). Neandethals also have moe obust distal aticulations than ealy moden humans but ae not diffeent in this aspect fom ecent moden humans (Figue 5-54). Ealy moden humans have longe ulnae with elatively lage poximal heads than those of ecent moden humans but ae simila in othe aspects of thei mophology (Appendix 48) (Figue 5-45; Figue 5-46). 305

337 Table 5-22 Desciptives of palaeogoup and the univaiate measuements of the adius. Max_ Length Olecshaftatio MidShaftShape Radial Notch Suface atio TochNotchOi OlecOient angle CoOleRatio BachRatio onato cest length Robusticity at 50% Robusticity at 25% Robust dist atic Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human Neandethal Ealy moden human Recent moden human N Mean S.D

338 Table 5-23 ANOVA esults fo palaeogoup and univaiate measuements of the ulna. d.f.=2 Max_ Length Headshaftation MidShaftShape Radial Notch Suface atio TochNotchOi headoient angle CoOleRation BachRatio Size ponato cest el. length Robusticity at 50% Robusticity at 25% Robust dist atic *=significant at α=0.05 F Sig * <0.001* 0.020* 0.019* 0.028* 0.003* 0.002* <0.001* * * Figue 5-45 Maximum ulna length fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 307

339 Figue 5-46 Olecanon-shaft atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). Figue 5-47 Midshaft shape atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 308

340 Figue 5-48 Radial notch suface aea fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). Figue 5-49 Tochlea notch oientation fo Neandethals, ealy and ecent moden humans. A lowe value is a moe anteioly facing tochlea notch compae to a poximo-anteioly facing one. Mean and 95% confidence inteval (whiskes). 309

341 Figue 5-50 Olecanon oientation fo Neandethals, ealy and ecent moden humans. A lowe value is a moe anteioly facing tochlea notch compae to a poximo-anteioly facing one. Mean and 95% confidence inteval (whiskes). Figue 5-51 Coonoid-olecanon atio fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 310

342 Figue 5-52 Bachial insetion atio fo Neandethals, ealy and ecent moden humans. A highe value means a elatively lowe insetion. Mean and 95% confidence inteval (whiskes). Figue 5-53 Ulna obusticity at 50% shaft level fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 311

343 Figue 5-54 Robusticity of the head of the ulna aea fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes) Disciminant function analysis A DFA with coss-validation using all adius Cs used in the analyses above and univaiate measuements of the adius was used to sepaate Neandethals, ealy and ecent moden humans (Figue 5-55). Function 1 sepaates best between Neandethals and all moden humans, wheeas function 2 sepaates ealy moden humans fom ecent moden humans. Function 1 eflects by (odeed accoding to deceasing coelation between the vaiable and the function) mcuveallc1 (medial cuvatue), head shape atio, lcuveallc3, lcuveallc2, position of the adial tubeosity, lcuveallc1 (lateal cuvatue), neck-length atio, and poximal and distal aticulation size atio. Function 2 eflects the length of the adius, midshaft shape, neck-shaft angle, head obusticity, mcuveallc2, mcuveallc3, EpiAllC1 and midshaft obusticity (Table 5-24). Fo all thee goups (Neandethals, ealy and ecent humans) with vey uneven sample sizes, the expected popotion of coect andom classification based on sample sizes is ~85%. The DFA with coss-validation using all Cs fo the adius included in the above analyses and univaiate 312

344 measuements was able to classify Neandethals and ecent moden humans with 50% (7 out of 14) of Neandethals and 83% (289 out of 348) of moden humans coectly classified. The ealy moden humans wee classified coectly in 50 % (7 out of 14) of cases, 14.3% (2 out of 14) wee classified as Neandethals. This gives an oveall coect classification of 80.6%. Function 2 Function 1 Figue 5-55 Disciminant Function 1 and 2 fo fo Neandethals, ealy and ecent moden humans. 313

345 Table 5-24 Disciminant function coefficients adius. Function 1 McuAllC1 DosalST HeadShapeRatio LcuAllC3 LcuAllC2 osradtubml distatshaftsizeratio LcuAllC1 NeckLengthRatio McuAllC3 CondAllC2 Canonical R= Λ = <0.001 Function 2 Max_ Length LatealST midshaftshaperation Headobusticity neck-shaft angle McuAllC2 HeadShaftSizeRatio CondAllC1 Midshaftobusticity Canonical R= Λ = <0.001 A DFA using all ulna Cs and univaiate measuements used in the analyses above was used to distinguish between Neandethals, ealy and ecent moden humans (Figue 5-56). Function 1 sepaates best between Neandethals and moden humans in geneal, wheeas function 2 sepaates ealy moden humans fom ecent moden humans. Function 1 eflects (odeed accoding to deceasing coelation between the vaiable and the function) poxallc3 (diection of the tochlea notch), neck-shaft angle, suface aea of the adial notch, obusticity at 25% and negatively by bachial tubeosity atio, pcuveallc2, and obusticity at 50%. Function 2 eflects the length of the ulna, the size of the head, obusticity of the distal aticulation, coonoid-olecanon atio, oxallc2, pcuveallc1, poxallc1, ponato cest size, pcuveallc3, and pcuveallc4 (Table 5-25). Fo these thee populations with vey uneven sample sizes, the expected popotion of coect andom classification is ~83%. The DFA with coss-validation using all Cs fo the ulna included in the above analyses and univaiate measuements was able to classify 61.5% (9 out of 314

346 15) of Neandethals and 98.5% (356 out of 361) of ecent moden humans coectly. Ealy moden humans had low classification success: 94.7% (14 out of 15) was classified as ecent moden human, 5.3% (1 out of 15) wee classified as Neandethals. This gives an oveall coect classification of 92.3%. Funtion 2 Function 1 Figue 5-56 Disciminant Function 1 and 2 fo Neandethals, ealy and ecent moden humans. Mean and 95% confidence inteval (whiskes). 315

347 Table 5-25 Disciminant function coefficients - ulna Function 1 oxallc3 BachRatio pcuveallc2 Robusticity at 50% headoient angle Radi Notch Suf atio MidShaftShape TochNotchOi Robusticity at 25% Canonical R= Λ = <0.001 Function 2 Max_ Length Headshaftation Robust dist atic CoOleRation oxallc2 pcuveallc1 oxallc1 pon. c. Length atio pcuveallc4 pcuveallc3 Canonical R= Λ = < Summay Neandethals have moe cuved adii (medial and lateal) and a moe sinusoidally shaped shaft than moden humans, and ealy moden humans ae simila to ecent moden humans. Neandethals also have an apex of cuvatue at midshaft, wheeas that of moden humans is located poximally. The Neandethals ae not diffeent in anteoposteio sinusoidal shape fom moden humans. Neandethals ae chaacteised by a pooly pojected styloid pocess and less mediolateal cuvatue of the ulna shaft. Neandethals have the most anteio facing tochlea notch. Ealy moden humans have an intemediate and moden humans have the most poximoanteio tochlea notch. Neandethals have a lage poximal ulna with a small neckshaft angle, a low bachial tubeosity, and highe midshaft obusiticity of the ulna than ecent moden humans. 316

348 CHATER DISCUSSION AND CONCLUSION Discussion The goal of this eseach was to investigate the diffeences and similaities between Neandethals and moden humans in long bone cuvatue. Moe specifically, this study tested hypotheses to explain vaiation in cuvatue among pesent-day and Holocene populations of humans, and applied these esults to the intepetation of Neandethals. Since thee was elatively little infomation available about long bone cuvatue in moden humans, this study examined geogaphically, tempoally and behavioually divese moden human samples in ode to evaluate coelates of longitudinal long bone cuvatue such as body size, climate, habitual behaviou, and mobility. The femu and adius wee chosen because they have been descibed as highly cuved in Neandethals (Ried, 1924; atte, 1955; Chuchill, 1998). The ulna was included because the shape of the adius can only be fully undestood if its inteaction with the ulna is also investigated. Limitations of pio eseach on cuvatue may have been due to the difficulty of accuate quantification which is appaent fom the inconsistency in techniques epoted in the liteatue (Ried, 1924; Genna, 1930; Stewat, 1962; Walensky, 1962, 1965; Gilbet, 1975, 1976; Tudell, 1999; Shackelfod and Tinkaus, 2002). Theefoe, it was necessay to find a method to quantify the patten of longitudinal bone cuvatue that would accuately epesent the thee-dimensional aspect of the diaphyseal suface and eliminate effects of scale. Thee dimensional geometic mophometics have fequently been used in canial eseach (fo an oveview see Slice, 2005), but its application to postcania is ae, and this method has not peviously been used to quantify long bone cuvatue in humans and thei close elatives. Hee, the method was successfully tested fo inta-obseve eo and shown to distinguish among populations moe effectively than taditional methods, such as diect measuement of subtense. In Chapte 4, the esults of the cuvatue analyses fo the femu, ulna and adius wee pesented fo the ecent human samples. These esults suggest that thee ae pattens within long bone cuvatue but that these ae diffeent fo the uppe and lowe limb. Femoal cuvatue is elated to habitual activity pattens. The highest levels of cuvatue fo the femu wee found in 317

349 populations with the highest activity levels. Femoal cuvatue follows diffeent tends fom obusticity and is not necessaily a esponse to the same loading egime (Ruff et al., 1993; Ruff et al., 1994; Tinkaus et al., 1994; Tinkaus et al., 1999a; eason, 2000b; Ruff and Tinkaus, 2000; Shackelfod and Tinkaus, 2002; Stock, 2002; Stock and feiffe, 2004; Stock, 2006; Calson et al., 2007; Shackelfod, 2007). Fo the femu, which is loaded poximodistally, cuvatue lowes bending stess by tanslating bending stess to axial compession (Fost, 1967; Hall, 2004), and cuvatue may be a compomise between bone stength and pedictability of bending stains and mateial failue (Lanyon, 1980, 1987; Betam and Biewene, 1988). Because femoal cuvatue is unelated to climate (latitude in this analysis), it may ultimately be a bette indicato of activity levels than coss-sectional measues of long bone obusticity. In contast, vaiation in cuvatue of the adius and ulna is influenced by climate. Individuals fom colde climates tend to have moe cuved and moe sinusoidal adii. Consistent with Begmann s (1847) and Allen s (1877) ules on body size and popotions, human populations fom colde climates have shotened distal limb segments, and it is thought that these diffeences ae genetic adaptations athe than epigenetic outcomes (Begmann, 1847; Allen, 1877; Y'Edynak, 1976; Eveleth and Tanne, 1990; Ruff et al., 1994; eason, 2000b; Van Andel, 2003; Weave, 2003; Ruff et al., 2005). The esults fo the ecent moden human sample suggest that cuvatue of the foeam bones is also a consequence of long-tem exposue to cold climate conditions athe than as a esult of habitual behaviou. This cuvatue is aguably not an adaptation in itself, but a consequence of educed elative foeam length in cold-adapted populations. In ode to optimise stength of the foeam despite its shote length, cuvatue may seve to maintain full function of the ponation and supination muscles, peseve inteosseous suface aea and facilitate muscle packing by allowing fo the position of slende attachments close to the joints while poviding adequate space fo the muscle bellies in the midshaft egion (Lanyon and Boun, 1979; Lanyon, 1980) (see Chapte 4). In Chapte 5, vaiation between Neandethals, ealy and ecent moden humans was evaluated (Objective 2). Neandethals ae distinct fom both ealy and ecent moden humans and exhibit a highe degee of anteio femoal cuvatue and a highe degee of lateal and medial cuvatue of the adius. Thee ae no diffeences in anteoposteio sinusoidal shaft shape of the ulna (posteio subtense) but Neandethals ae less mediolateally sinusoidal than ealy and ecent moden humans. 318

350 Based on pevious eseach, Neandethals ae thought to show evidence of cold-adaption in thei femoa, adii and ulnae. Fo the femu, Neandethals have extemely lage femoal heads and knees (distal ends) which ae consistent with thei cold-adapted body popotions and elatively lage body size (Tinkaus, 1981; Ruff, 1991; Chuchill, 1998; Weave, 2003). Both the adius and ulna ae elatively shot and also have lage aticulations. This shows that Neandethals confom to Begmann s (1847) and Allen s (1877) ules and that Neandethals fall at the cold end of the distibution, moe exteme than moden human populations. Some have suggested that the effects of, fo example, foeshotening of the distal extemities is not a heat consevation mechanism educing suface aea, but instead is the effect of the cooling of distal segments of the limbs and slowing of the metabolism and gowth of the peipheal tissues (Steegmann J. et al., 2002). Howeve, body shape manifests itself in ealy fetal life (Waen, 1998; Holliday, 2000) and does not appea to change with the secula tend in moden humans that affects body size and statue. Theefoe, it is likely to be genetically contolled (Katzmazyk and Leonad, 1998). The exteme cold-adapted physique of Neandethals can also be explained by thei lifestyle (Chuchill, 1998). Although Neandethals would have needed additional potection fom the cold in ode to suvive in Euope (Aiello and Wheele, 2003), the sevee physical stess of living in the Late leistocene cold Euopean and Westen Asian climate with simple technology may be sufficient to explain thei hype-pola body fom (Chuchill, 1998). The Mousteian (with which most Neandethals ae associated) does not show much evidence fo cultual buffeing against the cold. In contast, the Uppe alaeolithic tool industies ae typified by the fist solid evidence fo the systematic constuction of complex heaths suitable fo intensifying and containing heat (James et al., 1989; Stine, 1993; Tinkaus and Shipman, 1993 but see Heny et al., 2004). unches o awls and the subsequent appeaance of needles epesent advances in making tools fo binding hide togethe fo clothing (Tinkaus, 1981; Holliday, 1997; Holliday, 1999; Weave and Steudel-Numbes, 2005 and aticles in Mellas and Stinge, 1989). The lack of such technological advances in themal potection in Neandethals may explain the selective pessues on them while thei pesence in moden achaeological assemblages may point to the eduction in those selective pessues in moden humans inhabiting simila climates (Rak, 1990; Tinkaus et al., 1998a; Holliday, 2000; Chuchill, 2001; Niewoehne, 2001; Shea, 2003). 319

351 Although the Neandethal femu shows some climatic adaptations (Tinkaus, 1981; Ruff, 1991; Chuchill, 1998; Weave, 2003), the esults fom the ecent human analyses indicate that thee is no effect of climate on the cuvatue of the femu (Walensky, 1965; Gilbet, 1976). In addition, cuvatue is not coelated to femoal tosion. This is consistent with femoal cuvatue being unelated to climate because, if femoal cuvatue was a consequence of the wide coldadapted Neandethals hip, it would be coelated with tosion as was suggested by Weave (2003). The cuvatue of the adius and oveall mophology of the ulna in Neandethals shows good coespondence with the climate data. This climatic vaiation is confimed also in the distibution of the medial adial cuvatue: fossils fom Neandethal populations in colde climates (Spy, Le Moustie, La Quina, La Feassie) have a highe degee of cuvatue than the Middle Easten fossils (Shanida and Kebaa). As discussed above, the elationship between cuvatue and climate can be explained as a consequence of the shotening of the foeam. Othe chaacteistics in the foeam that ae coelated with climate in ecent moden humans ae a moe sinusoidal adial shaft, shotening of the ulna neck (distance between 80% level of the ulna and the tip of the coonoid), a poximo-anteioly facing tochlea notch, inceased distal aticulation size, a lage ulna head elative to shaft, lage adial notch suface aea and a elatively lowe insetion of the bachial muscle. These featues indicate that the absolute dimensions of the head and aticulations of the ulna and adius emain elatively lage fo the length of the shaft. Also, foeshotening of the foeam in esponse to cold climatic conditions is not a scaling down of the whole bone but athe a eduction in shaft length. Shotening the diaphysis educes the suface aea fo muscle insetions and may affect leve advantage and contaction function (which is affected by muscle fibe size) of seveal am and hand muscles, such as the ponato tees (Hall, 2004). Theefoe, cuvatue may be a means of maintaining full function and foce despite a eduction in length. By inceasing the cuvatue of the adius and adopting a moe sinusoidal shaft shape diaphyseal length is maximised. The esults fo the Neandethal ulna and adius show that Neandethals have all the above mentioned cold climate featues and expess them to a moe exteme degee. Neandethals have the highest degee of lateal cuvatue of the adius, elatively the lagest ulna head, shotest ulna neck (distance between 80% of the shaft and the tip of the coonoid), the most 320

352 anteioly facing tochlea notch and the most infeio bachial tubeosity. The emegence of moden humans saw a ponounced eduction in the muscula hypetophy of the uppe limb (Tinkaus, 1986) and a eduction in the size of the muscle insetions on the am and hand skeleton (ponato quadatus on the ulna, the flexo pollicis longus and the opponens muscles on the capals and distal phalanges) (Tinkaus, 1983a). The eduction in muscle size in moden humans may also explain the lowe degee of cuvatue in moden-day Actic populations compaed to Neandethals (Lanyon and Boun, 1979). The Neandethal adius shows some distinct featues such as a moe medially placed adial tubeosity compaed to moden humans. It has been suggested that this position of the adial tubeosity is a consequence of the use of the foeam in flexion (Tinkaus and Chuchill, 1988). In the pesent study it was pedicted that if cuvatue of the adius is a esult of the habitual use of the am in that position and the associated inceased stain of the foeam, that thee would be a coelation between the position of the adial tubeosity, the neck-length of the adius and cuvatue. Neandethals do have a moe medially oiented adial tubeosity than do moden humans, but thee is no coelation with neck length o with cuvatue. Also, in moden humans a moe medially oiented adial tubeosity was associated with low activity levels. The ecent moden human analyses suggested that femoal cuvatue is a plastic featue that esponds to loading of the femu duing activity. Confiming the hypothesis by Shackelfod and Tinkaus (2002), populations with high activity levels have a high degee of femoal cuvatue. This was evident also in the elationship between activity levels and obusticity at diffeent points along the shaft. It is not supising, then, that thee is a elationship between cuvatue and obusticity in moden humans. The coelation between coss-sectional anteoposteio obusticity and activity levels was hypothesised to be the esult of epetitive loading on the lowe limb duing subsistence stategy-elated teestial mobility (Ruff, 1987; 1994; Lasen et al., 1995; Holt, 2003; Stock and feiffe, 2004), and this hypothesis is suppoted by the stength ciculaity indices at the femoal midshaft and thei stong coespondence with teestial mobility (Stock, 2006). Because of the coelation between subsistence-elated activity and cuvatue in ecent moden humans, the pediction was that Neandethals, being huntegathees, would have a high degee of femoal cuvatue. Moeove, thei cuvatue should be compaable to that of ealy moden humans because the two goups had boadly simila lifestyles (Tinkaus et al., 1989). 321

353 Ealy moden humans and Neandethals most likely did not diffe in thei subsistence stategies and wee both hunting and scavenging (Liebeman, 1989; Ba-Yosef, 2004; eason et al., 2006). Faunal assemblages fom occupation and butchey sites shows that both goups had ealy access to the animals and cut-mak pattens indicate a pimay eliance on hunting athe than scavenging (Speth and Tchenov, 1998). Tinkaus and Zimmeman (1982) and Klein (2003) have agued that Middle Stone Age people wee less adept huntes because they only hunted a few of the available species and that Neandethals show a high incidence of skeletal tauma because of the isk involved in close ange hunting (Tinkaus and Zimmeman, 1982; Klein, 2003). Recent investigations of faunal assemblages have shown that some Neandethal sites may be dominated by a single pey species, but this is also documented among some moden huntegathee societies (Maean and Assefa, 1999; Maean and Assefa, 2005). The eliance on meat fo Neandethals and ealy moden humans living in tempeate and cold egions such as Euope and Westen Asia was impotant fo suvival. Ealy Euopeans must have elied on fequent meat acquisition fo thei diet as it is likely that plant foods would have been unavailable fo consumption duing pats of the yea. This is confimed in stable-isotope analyses fom sites such as Vindija Cave, Coatia; Scladina, Spy and Engis in Belgium and Maillac and Saint-Césaie in Fance (Fizet et al., 1995; Richads et al., 2000; Bocheens et al., 2001; Richads et al., 2001; Ducke and Bocheens, 2004; Bocheens et al., 2005). Maean (1999) agued that the Middle alaeolithic Neandethals may not have been less adept huntes than thei Middle Stone Age moden human contempoaies but, instead, might have been less adept at using and pocessing cacasses in ode to ende highe caloic yields, such as fat endeing and stoage, which put them at a subtle disadvantage in compaison to moden humans. These disadvantages wee not only the lowe caloic intake pe pey animal, but also the inceased pesonal isk because of moe fequent hunting (Maean and Assefa, 1999). This low etun on time expended may have esulted in modeately highe activity and mobility levels in Neandethals compaed to ealy anatomically moden humans. Similaities in lifestyle and subsistence patten between Neandethals and the ealiest moden humans is also appaent in the achaeological ecod, whee simila species of lage animals ae found in both Neandethal and ealy moden human deposits. Neandethals wee effective huntes (Speth and Tchenov, 1998) and some conside them a top pedato in the envionment in which they lived (Bocheens et al., 2005). They also hunted a given egion fo a longe peiod 322

354 of time than moden humans who wee moe seasonally mobile (Liebeman, 1989). Although thee is some vaiation, oveall, Neandethals and ealy moden humans wee likely vey simila in tems of mobility, esouce acquisition and oveall wokload, and this is appaent in thei postcanial anatomy (Liebeman, 1989). When coected fo size and body popotions, Neandethals have lowe limb bones that wee simila in coss-sectional stength to those of moden humans (Tinkaus et al., 1989). This is also eflected in the esults on obusticity pesented hee, which showed no significant diffeences between obusticity levels of the shaft between Neandethals and ealy moden humans. In degee of femoal cuvatue, howeve, and contay to the hypothesis of Shackelfod and Tinkaus (2002), Neandethals show a significantly highe degee of cuvatue and a lowe apex of cuvatue compaed to both ealy and ecent moden humans. This diffeence suggests that Neandethals had much highe activity levels, in contast to what is suggested by the obusticity esults (Tinkaus et al., 1989). The compaatively small ange of vaiation in femoal cuvatue in Neandethals compaed to ealy and ecent moden humans (and in paticula compaed to the ange of vaiation of the high activity goup) suggests that Neandethals had a smalle ange of subsistence behavious than moden humans and that this behaviou involved high activity levels. The cuvatue of the adius is a eflection of climate and the vaiation among Neandethals and ealy moden humans is vey wide compaed to that of ecent moden humans. Also Neandethal adii fom the Levant tend to be less cuved than those fom Nothwest Euope. Most ealy moden humans fall within the ange of ecent moden humans, but Skhul IV and Qafzeh 9 fall outside. The sites of Skhul and Qafzeh ae the ealiest moden human sites outside of Afica, and it has been suggested that the individuals fom these sites wee not yet fully moden human (McCown and Keith, 1939b, 1939a; Aensbug and Belfe-Cohen, 1998; Kame et al., 2001; Rak, 2002). The ealy moden humans fom Skhul and Qafzeh also pe-date the pesence of Neandethals in the egion, and some have suggested that the distinctiveness of Neandethal vesus moden humans in the Levant may not be as clea as in othe places, and the ovelap in mophology may be explainable by admixtue between the two goups (Kame et al., 2001). This may also explain the highe degee of adial cuvatue obseved in those two individuals compaed to the est of the goup. In light of the ecent genetic evidence showing that Neandethals did not contibute to the moden human gene pool (Caamelli et al., 2003; Ovchinnikov and Goodwin, 2003; Geen et al., 323

355 2006) the diffeences in cuvatue of the Skhul and Qafzeh could be explained by the vey ealy date fo these individuals if thee was evidence fo inceased musculaity elative to moe ecent moden humans. This has been contadicted by studies on the humeus and hand bones which showed that the ealy Nea Easten moden humans wee moe gacile than Middle Stone Age and late Uppe alaeolithic moden humans and wee thus somewhat of an anomaly (Tinkaus and Chuchill, 1999; Niewoehne, 2001). The evolutionay significance of long bone cuvatue fo hominins moe geneally has not been investigated. The femu and adius of goillas and chimpanzees ae moe cuved than those of moden humans (Matin and Salle, 1959), and long bone cuvatue in pimates is known to scale positively with body weight (Swatz, 1990). In humans, thee is no coelation between body size and cuvatue, but the vaiation in moden human long bone cuvatue shows that, despite not being allometically scaled, its plasticity was etained thoughout human evolution and cuvatue should theefoe be consideed a selectively adaptive featue. With the shift to bipedal walking in hominins, weight distibution and muscle oganisation of the femu has changed, and the uppe limb lost its locomoto function. The functional significance of long bone cuvatue in ealie hominins has not been commented on, but it is possible to examine some hominin casts and published photogaphs. The Homo eectus Naiokotome femu is elatively staight but the Homo sp. KNM-ER 1481 shows a maked degee of femoal cuvatue despite having a elatively gacile shaft. hotogaphs of elatively complete femoa fom othe membes of the genus Homo, such as those fom Atapueca and Dmanisi, have only been published in anteio view, so it is impossible to comment on the degee of femoal cuvatue (Lodkipanidze et al., 2007). Radii ae pooly epesented in the fossil ecod. The fagmentay adius fom OH 62 and a fagment fom KNM-ER 3735 indicate a modeately cuved adius fo H. habilis (Haeusle and McHeny, 2004), and a adial fagment fom Atapueca suggests a low degee of adial cuvatue fo H. antecesso. Although complete and well dated postcanial fossils ae elatively ae, the use of 3D geometic mophometics on both complete and patial fossil specimens and a compaison with the Afican apes should povide sufficient data fo investigating futhe the evolutionay significance of long bone cuvatue in ealie hominins. 324

356 6.2. Conclusion The evidence pesented hee suppots the hypothesis that femoal cuvatue is a bone esponse to stesses and stains duing habitual behaviou and shows good coespondence with measues of extenal obusticity. opulations with high activity levels have a highe degee of anteio femoal cuvatue and a moe distal apex of cuvatue than populations with low and modeate activity levels. Within populations with high activity levels, males have moe cuved femoa than females. This is not due to sexual dimophism in body size o sex diffeences in bone modelling and emodelling as thee is no sex diffeence in goups who have less sexual division of labou and cuvatue is not coelated to body size. Of the high activity subsistence stategies the aquatic foages, with low levels of teestial activity, ae the least cuved, and the pastoalists, with high levels of teestial mobility, ae the most cuved. Biomechanically, inceased femoal cuvatue seves to geneate physiologically beneficial stains, facilitates muscle packing and inceases the pedictability of mateial failue. Lateal cuvatue of the adius, mediolateal cuvatue of the ulna and oveall foeam bone shape ae coelated with climate and ae poo pedictos of habitual behaviou. Howeve, the aquatic foages wee distinct in having a poximal development on the inteosseous cest and high neck-shaft angles which may eflect thei use of watecaft. Cuvatue of the adius and ulna is likely a consequence of the foeshotening of the foeam in cold-adapted populations. The esults suggest that this foeshotening is a eduction in length of the diaphysis while maintaining elatively lage epiphyses and athe than an oveall downscaling of the bone. Inceased foeam bone cuvatue aids in maintaining the tendon insetions close to the joints while facilitating muscle packing, and etaining inteosseous space, muscle length and function and maximising diaphyseal length. Neandethals and ealy moden humans had boadly simila hunte-gathee lifestyles, and thei postcanial skeleton was likely subject to the same stesses as moden humans. Neandethals show a high degee of femoal cuvatue, eflecting thei active lifestyles, and a high degee of adial cuvatue, eflecting thei cold-adapted body fom. Ealy moden humans display a high degee of femoal cuvatue but, contay to Neandethals, one that is well within the ange of vaiation of moden humans. Ealy moden humans, except fo Skhul and Qafzeh, show a ange 325

357 of vaiation of adial cuvatue that falls within the ange of ecent moden humans. Neandethals fall above the human ange of cuvatue, although thee is some ovelap. Neandethals also show a numbe of diffeences in the shape of the ulna (e.g., moe anteioly facing tochlea notch and shote ulna neck). Although thee may have been some vaiation in the specific subsistence-elated activities they pefomed, thee is a widely held view that Neandethals and ealy moden humans had simila lifestyles and activity levels. Theefoe, the highe degee of femoal cuvatue in Neandethals cannot be explained by behaviou alone. Fom a taxonomic and phylogenetic pespective, Neandethals ae distinct in thei expession of cuvatue compaed to moden humans, but it emains to be investigated whethe the low degee of cuvatue is a deived ecent human tait, o whethe a maked degee of cuvatue is an autapomophy of Neandethals. It has also been suggested that cetain diffeences in mophology between Neandethals and ecent moden humans ae the esult of behaviou duing ontogeny (Tinkaus, 1993), but the esults pesented in this study would suggest that only femoal cuvatue may be affected by individual ontogeny. In ode to investigate this futhe it is necessay to expand this study to an ontogenetic sample. Numeous studies have emphasised the unusual featues of Neandethals and highlighted diffeences between them and us. As othe studies have done (see Tinkaus, 1975; Tinkaus and Villemeu, 1991; Chuchill, 1998, 2001; Weave, 2003; eason et al., 2006), this study used models based on vaiation in Homo sapiens to discuss postcanial mophology in the context of evolutionay biology and adaptive histoy of moden humans. Neandethals had active lifestyles and wee adapted to life in the cold climate of Euope and Westen Asia, but they wee diffeent fom cold-adapted moden humans and equally active ealy moden humans. Even when only the available evidence fom ealie membes of the genus Homo is taken into account, Neandethal femoal and adial mophology appeas to be distinct, especially when it is consideed in combination with the est of Neandethal postcanial chaactes. This distinctiveness can most easily be explained by the isolation of the Neandethals duing an extended peiod of time fom the moden human lineage. 326

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385 AENDIX Appendix 1 Landmaks and measuements fo the femu N Measuement and landmak Subtochanteic mediolateal diamete (Matin n 9) M 80% L 80% Midshaft mediolateal diamete (Matin n 8) M 50% L 50% Subpilastic mediolateal diamete M 25% L 25% Subtochanteic anteoposteio diamete (Matin n 11) A 80% 80% Mid-shaft anteoposteio diamete (Matin n 10) A 50% 50% Subpilastic anteoposteio diamete Desciption Medio-lateal diamete taken at the 80% level. The 0% shaft level is defined as the most infeio edge of the medial condyle; the 100% is the most supeio point of the head of the femu. Most medial point at 80% level. Most lateal point at 80% level Medio-lateal diamete taken at the 50% level. The 0% shaft level is defined as the most infeio edge of the medial condyle; the 100% is the most supeio point of the head of the femu Most medial point at 50% level. Most lateal point at 50% level. Medio-lateal diamete taken at the 25% level. The 0% shaft level is defined as the most infeio edge of the medial condyle; the 100% is the most supeio point of the head of the femu Most medial point at 25% level. Most lateal point at 25% level. Anteo-posteio diamete taken at the 80% level. The 0% shaft level is defined as the most infeio edge of the medial condyle; the 100% is the most supeio point of the head of the femu Most anteio point at 80% level. Most posteio point at 80% level. Anteo-posteio diamete taken at the 50% level. The 0% shaft level is defined as the most infeio edge of the medial condyle; the 100% is the most supeio point of the head of the femu Most anteio point at 50% level. Most posteio point at 50% level. Anteo-posteio diamete taken at the 25% level The 0% shaft level is defined as the most infeio edge of the medial condyle; the 100% is the most supeio point of the head of the femu

386 A 25% 25% Femu length (Matin n 1) FEML1 FEML2 Length of the head-neck axis (Matin N 14) HNAX1 HNAX2 Head diamete (Matin N 18 and 19) HDIA1s HDIA2i HDIA3p HDIA4a Neck-shaft angle (Matin N 29) HNAX1 NSAG2 NSAG3 Tosion (Matin n 28) HNAX1 TORS2 TORS3 355 TORS4 Most anteio point at 25% level. Most posteio point at 25% level. Maximum length measued along the biomechanical axis.(biomech axis: whee the most supeio point of the head of the femu and the most lateal point of the geate tochante descibe a 90 angle, the pependicula line down fom the most supeio point of the head to the most infeio point on the medial condyle). The most supeio point of the head measued along the biomechanical axis. The most infeio point on the medial condyle measued along the biomechanical axis. Length of the axis fom the most medial point of the head to the middle of the intetochanteic line. Most medial point of the head Middle of the intetochanteic line Maximum diamete of the femoal head on the edge of the aticula suface Most supeio point on a line descibing the maximum supe-infeio diamete Most infeio point on a line descibing the maximum supeo-infeio diamete Most posteio point on a line descibing the maximum mediolateal diamete Most anteio point on a line descibing the maximum mediolateal diamete Also collo-diaphyseal angle. Matin n 29. The angle descibed by the shaft-axis (going though the middle of the shaft) and the neck-axis (going though the middle of the neck) Most medial point of the head oint whee the neck axis intesects with the axis though the middle of the shaft Located on the supeio edge of patella suface midway between medial and lateal bodes of supeio potion of the patella suface. Also lies on line passing though middle of axis of the distal shaft. The angle of femoal tosion is the angle made by the axis of the femoal neck with the tangent of the posteio sufac of the femoal condyles. Most medial point of the head The most posteioly pojecting point of the medial condyle. The point whee the condyle would touch a suface if it wee hoizontally placed on a suface. The most posteioly pojecting point of the lateal condyle. The point whee the condyle would touch a suface if it wee hoizontally placed on a suface. Most lateal point on the geate tochante on the neck axis

387 12 Middle of the insetion aea fo gluteus minimus (Weave n 3) GMIN 13 Middle of the insetion aea fo gluteus medius (Weave n 4) GMED 14 Tip of the lesse tochante (Weave n 5) LSTR Tip of the adducto tubecle (Weave n 8) Located on the anteo-infeio suface of the geate tochante, just medial to the lateal bode, in the cente of the oval insetion aea fo gluteus minimus. The insetion aea may extend as a thinne stip supeioly and medially, but ecod the point in the cente of the insetion. Cente of the oval insetion aea fo gluteus minimus on the anteo-infeio suface of the geate tochante, medial to the lateal bode Located on the posteo-supeio suface of the geate tochante, in the cente of the oval insetion aea fo the gluteus medius. The insetion aea extends as a thinne stip infeioly and anteioly, but ecod the point in the cente of the insetion. Cente of the oval insetion aea fo the gluteus medius located on the posteo-supeio suface of the geate tochante Whee the lesse tochante pojects maximally (local maximum of a cuved suface) The tip whee the lesse tochante pojects maximally Located whee the adducto tubecle pojects maximally (local maximum of a cuved suface) 356 ADTB Midpoint of the anteosupeio edge of the patella suface of the distal femu (Weave n 9) NSAG3 Tip located whee the adducto tubecle pojects maximally Located on the supeio edge of the patella suface midway between the medial and lateal bodes of the supeio potion of the patella suface. This point also lies on a line that passes though the middle of the axis of the distal femoal shaft. Midpoint of the medial edge of the infeio suface of the medial condyle (Weave n 10) MCMDi Midpoint of the lateal edge of the infeio suface of the infeio suface of the lateal Midpoint, fom an infeio view, of the medial edge of the infeio suface of the medial condyle. Located on the sup edge of patella suface midway between medial and lateal bodes of supeio potion of the patella suface. Also lies on line passing though middle of axis of the distal shaft. Midpoint, fom an infeio view, of the medial edge of the infeio suface of the medial condyle. The midpoint, fom an anteio view, of the lateal edge of the infeio suface of the lateal condyle. Thee is usually a slight notch o depession at this point. oints 27 and 28 should connect to fom a line that is hoizontal when the femu is held in anatomical position. oints 27 and 28 usually fall just anteio to the anteio edge of the intecondyla notch.

388 condyle (Weave n 11) LCMDi Midpoint of the medial edge of the posteio suface of the medial condyle (Weave n 12) MCMDp Midpoint of the lateal edge of the posteio suface of the lateal condyle (Weave n 13) LCMDp Maximum condyla width (Matin n 21) MLMDM1 MLMDL2 Most supeio pojection of the patella suface ROJ1 ROJ2 Cuvatue Midpoint, fom an infeio view, of the lateal edge of the infeio suface of the lateal condyle. This point is defined as the midpoint, fom a posteio view, of the medial edge of the posteio suface of the medial condyle. Midpoint, fom a posteio view, of the medial edge of the posteio suface of the medial condyle. This point is defined as the midpoint, fom a posteio view, of the lateal edge of the posteio suface of the lateal condyle. Thee is usually a slight notch o depession at point 30. oint 29 and 30 should connect to fom a line that is a fontal plane when the femu is held in anatomical position. Midpoint, fom a posteio view, of the lateal edge of the posteio suface of the lateal condyle. The distance between the point whee the medial epicondyle pojects maximally (local maximum of a cuved suface) and the point whee the lateal epicondyle pojects maximally (local maximum of a cuved suface) The point whee the medial epicondyle pojects maximally (local maximum of a cuved suface) The point whee the lateal epicondyle pojects maximally (local maximum of a cuved suface) The points on a cuved suface whee the diection of the aticulation of the patella suface changes diection (fom lateal/medial to infeio) The point whee the medial condyla aticula suface pojects most anteioly The point whee the lateal condyla aticula suface pojects most anteioly Cuvatue of the femu along fou sides. osteio measued fom 80% level along the linea aspea down to the midpoint between the posteio medial and lateal patella suface. Anteio cuvatue measued fom the 80% level down to the midpoint on the most supeio edge of the patella suface. Medial cuvatue fom 80% level down to the adducto tubecle. Lateal cuvatue fom 80% level down to the LMXTB The point whee the lateal suface pojects maximally (opposite side adducto tubecle) MIDS The midpoint between the posteio medial and lateal patella suface CURV Semi-landmaks taken evey 5 mm along the posteio cuve of the femu. ACURV Semi-landmaks taken evey 5 mm along the anteio cuve of the femu. MCURV Semi-landmaks taken evey 5 mm along the medial cuve of the femu. LCURV Semi-landmaks taken evey 5 mm along the lateal cuve of the femu. Midshaft obusticity index A diamete 50% + ML diamete 50% / length *100

389 Head obusticity index Condyle diamete atio Neck length atio Subtochanteic atio Midshaft atio Subpilastic atio SI head diamete + A head diamete / length *100 Maximum condyla width/length *100 Neck length/length *100 A diamete 80% / ML diamete 80%*100 A diamete 50% / ML diamete 50%*100 A diamete 25% / ML diamete 25%*

390 Appendix 2 Landmak diagam femu (Afte ). 359

391 Appendix 3 Landmaks and measuements fo the adius N Measuement and landmak Maximal length (Matin n 1) RADL1 RADL2 80% mediolateal diamete (Matin 5a) M 80% L 80% 50% mediolateal diamete M 50% L 50% 25% mediolateal diamete M 25% L 25% 80% anteoposteio diamete (Matin 4a) A 80% 80% 50% anteoposteio diamete (Matin 5a) A 50% 50% 25% anteoposteio diamete A 25% Desciption Maximum length measued fom the most supeio point on the aticula suface on the head to the most distal point on the styloid pocess. The most supeio point on the aticula suface on the head The most distal point on the styloid pocess Medio-lateal diamete taken at the 80% level. The 0% shaft level is defined as the most infeio edge of the styloid pocess; the 100% is the most supeio point on the aticula suface on the head. Most medial point at 80% level. Most lateal point at 80% level. Medio-lateal diamete taken at the 50% level. The 0% shaft level is defined as the most infeio edge of the styloid pocess; the 100% is the most supeio point on the aticula suface on the head. Most medial point at 50% level. Most lateal point at 50% level. Medio-lateal diamete taken at the 25% level. The 0% shaft level is defined as the most infeio edge of the styloid pocess; the 100% is the most supeio point on the aticula suface on the head. Most medial point at 25% level. Most lateal point at 25% level. Anteo-posteio diamete taken at the 80% level. The 0% shaft level is defined as the most infeio edge of the styloid pocess; the 100% is the most supeio point on the aticula suface on the head. Most anteio point at 80% level. Most posteio point at 80% level. Anteo-posteio diamete taken at the 50% level. The 0% shaft level is defined as the most infeio edge of the styloid pocess; the 100% is the most supeio point on the aticula suface on the head. Most anteio point at 50% level. Most posteio point at 50% level. Anteo-posteio diamete taken at the 25% level. The 0% shaft level is defined as the most infeio edge of the styloid pocess; the 100% is the most supeio point on the aticula suface on the head. Most anteio point at 25% level. 360

392 8 9 25% Length of the head-neck axis (Matin 1a) HDII4a RADT Supeio head diamete (Matin n 4 (1)) HDIS1m HDIS2l HDIS3p HDIS4a 10 Infeio head diamete (Based on Matin n 4 (1) ) HDII1m HDII2l HDII3p HDII4a 11 Neck-shaft angle (Matin n 7) HDII4a NSAG2 A 80% Most posteio point at 25% level. Length of the axis fom the most supeio point of the head to the adial tubeosity. Most anteio point on a line descibing the maximum diamete on the most infeio edge of the head The tip whee the adial tubeosity pojects maximally Maximum diamete of the adial head on the edge of the aticula suface Most medial point on a line descibing the maximum mediolateal diamete on the most supeio edge of the head Most lateal point on a line descibing the maximum mediolateal diamete on the most supeio edge of the head Most posteio point on a line descibing the maximum anteoposteio diamete on the most supeio edge of the head Most anteio point on a line descibing the anteoposteio maximum diamete on the most supeio edge of the head Maximum diamete of the femoal head on the edge of the aticula suface 361 Most medial point on a line descibing the maximum mediolateal diamete on the most infeio edge of the head Most lateal point on a line descibing the maximum mediolateal diamete on the most infeio edge of the head Most posteio point on a line descibing the anteoposteio maximum diamete on the most infeio edge of the head Most anteio point on a line descibing the anteoposteio maximum diamete on the most infeio edge of the head Also collo-diaphyseal angle. Matin n 7. The angle descibed by the shaft-axis (going though the middle of the shaft) and the neck-axis (going though the middle of the neck) Most anteio point on a line descibing the maximum diamete on the most infeio edge of the head oint whee the most naow diamete of the neck intesects with the anteio neck axis though the middle of the shaft Most anteio point at 80% level. The 0% shaft level is defined as the most infeio edge of the styloid pocess; the 100% is the most supeio point on the aticula suface on the head.

393 The adial tubeosity RADT Middle of the distal adial aticula suface edge ARTSp The middle of the ulna notch ULNT Middle of the distal adial aticula suface edge ARTSa Dosal subtense (based on Matin 6b) Lateal subtense (based on Matin 6a) Cuvatue MCURV LCURV Midshaft obusticity Head obusticity Distal aticulation Size Ratio osition Radial Tubecle Neck Length Ratio (Matin n 1a/Matin n 1) Head Shape Ratio Midshaft Shape Ratio (Matin n 4a/ Matin n 5a) The most pojecting point on the adial tubeosity The tip whee the adial tubeosity pojects maximally The middle of the distal adial aticula suface edge on the posteio side. The middle of a cuved suface The middle of the distal adial aticula suface edge on the posteio side. The middle of a cuved suface The middle of the aticula suface on the medial side of the adial notch. The middle of the medial aticula suface on the ulna notch The middle of the distal adial aticula suface edge on the anteio side. The middle of a cuved suface The middle of the distal adial aticula suface edge on the anteio side. The middle of a cuved suface Maximum distance fom a chod connecting 80% and ARTSp and the posteio suface of the shaft. Maximum distance fom a chod connecting L80% and the most distal point on the styloid pocess (RADL2) and the posteio suface of the shaft. Cuvatue of the adius along two sides. Medial cuvatue fom 80% down to the middle of the ulna notch. Lateal cuvatue fom 80% level down to the tip of the styloid pocess. Semi-landmaks taken evey 5mm along the medial cuve of the adius Semi-landmaks taken evey 5mm along the lateal cuve of the adius anteoposteio midshaft diamete + mediolateal midshaft diamete/ maximum length *100 anteopostio head diamete+ mediolateal diamete/ maximum length *100 anteoposteio distal aticulation diamete + mediolateal distal aticulation diamete/ maximum length *100 the angle between a vecto connecting the most pojecting point on the adial tubeosity and the most medial point at the 80% level and the vecto unning though the most medial point at 50% and 80% (see diagam) Neck length/maximum length *100 anteopostio head diamete / mediolateal diamete *100 anteoposteio midshaft diamete / mediolateal midshaft diamete *

394 Appendix 4 Landmak and measuement diagams adius (Afte and ) 363

395 364

396 Appendix 5 Landmaks and measuements fo the ulna N Measuement and landmak Maximum length (Matin n 1) ULNL1 ULNL2 80% mediolateal diamete M 80% L 80% 50% mediolateal diamete M 50% L 50% 25% mediolateal diamete M 25% L 25% 80% anteoposteio diamete A 80% 80% 50% anteoposteio diamete A 50% 50% 25% anteoposteio diamete A 25% 25% Desciption Maximum length measued fom the most supeio point on the olecanon pocess to the most distal point on the aticula suface (not styloid pocess because of pesevation issues in achaeological samples) The most supeio point on the olecanon pocess The most distal point on the adial aticulation suface Medio-lateal diamete taken at the 80% level. The 0% shaft level is defined as the most distal point on the aticula suface; the 100% is the most supeio point on olecanon pocess Most medial point at 80% level. Most lateal point at 80% level. Medio-lateal diamete taken at the 50% level. The 0% shaft level is defined as the most distal point on the aticula suface; the 100% is the most supeio point on olecanon pocess Most medial point at 50% level. Most lateal point at 50% level. Medio-lateal diamete taken at the 25% level. The 0% shaft level is defined as the most distal point on the aticula suface; the 100% is the most supeio point on olecanon pocess Most medial point at 25% level. Most lateal point at 25% level. Anteo-posteio diamete taken at the 80% level. The 0% shaft level is defined as the most distal point on the aticula suface; the 100% is the most supeio point on olecanon pocess Most anteio point at 80% level. Most posteio point at 80% level. Anteo-posteio diamete taken at the 50% level. The 0% shaft level is defined as the most distal point on the aticula suface; the 100% is the most supeio point on olecanon pocess Most anteio point at 50% level. Most posteio point at 50% level. Anteo-posteio diamete taken at the 25% level. The 0% shaft level is defined as the most distal point on the aticula suface; the 100% is the most supeio point on olecanon pocess Most anteio point at 25% level. Most posteio point at 25% level.

397 onato quadutus cest RQC1 RQC2 oximal aticulation dimension OLT OLMXm OLMXl OLMXp TRWD1 TRWD2 CORR RADNm RADNa RADNp Distal aticulation diamete HDIAp HDIAa HDIAm HDIAl Styloid pocess STR Flexo digitoum sublimis FLXSm Bachialis insetion BRACH1s The dimensions of the ponato quadutus cest The most poximal point of the ponato quadutus cest The most distal point of the ponato quadutus cest The dimensions of the olecanon and coonoid pocess. The tip of the Olecanon pocess The most medial point on the olecanon pocess The most lateal point on the olecanon pocess The most posteio point on the olecanon pocess The most medial point on the tochlea notch along the minimum width line pependicula to the shaft axis The most lateal point on the tochlea notch along the minimum width line pependicula to the shaft axis The tip of the coonoid pocess The most medial point on the adial notch The most anteio notch on the adial notch The most infeio notch on the adial notch Anteoposteio and mediolateal diamete of the supeio edge of the distal aticulation The most posteio point on the supeio edge of the distal aticulation The most anteio point on the supeio edge of the distal aticulation The most medial point on the supeio edge of the distal aticulation The most lateal point on the supeio edge of the distal aticulation The tip of the styloid pocess The tip of the styloid pocess The most pojecting tip of the flexo digitoum sublimis The middle of the flexo digitoum sublimis The dimensions of the bachialis insetion The most supeio point of the bachialis insetion

398 BRACH2i BRACH3m Cuvatue CURV tochlea notch oientation (Matin n 15) Olecanon size (atte, 1955; Fishe, 1906 p. 227) osition bachialis (Solan, 1992) Head oientation (Matin, 15a) Head/shaft atio Coonoid Olecanon atio (Matin 7a and 8a) The most infeio point of the bachialis insetion The middle of the bachialis insetion osteio cuvatue measued fom 80% level down to the most posteio point on the adial aticulation. Semi-landmaks taken evey 5 mm along the posteio cuve of the ulna The angle between the vecto unning along the anteio suface and the vecto connecting the tip of the olecanon and coonoid (also joint-axis angle) The distance between the tip of the olecanon and the most posteio point on the poximal suface of the ulna (see diagam) The position of the bachialis tubeosity: Distance fom the poximal extemity to the most distal point of the bachialis tubeosity (see diagam) The angle at the olecanon when a tiangle is fomed between the 80% anteio suface, the tip of the olecanon and the coonoid (see diagam) Size of the head: olecanon size/length *100 (see diagam) Height olecanon/height coonoid*100 (see diagam) 367

399 Appendix 6 Landmak and measuement diagams ulna (Afte and ) 368

400 Appendix 7 Diagams fo measuements calculated fom the landmaks on the ulna (Afte and ) 369