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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">The Influence of Life History and Sexual Dimorphism on Entheseal Changes in Modern Humans and African Great Apes</title><author><name sortKey="Milella, Marco" sort="Milella, Marco" uniqKey="Milella M" first="Marco" last="Milella">Marco Milella</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">25251439</idno><idno type="pmc">4175998</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4175998</idno><idno type="RBID">PMC:4175998</idno><idno type="doi">10.1371/journal.pone.0107963</idno><date when="2014">2014</date><idno type="wicri:Area/Pmc/Corpus">000029</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000029</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">The Influence of Life History and Sexual Dimorphism on Entheseal Changes in Modern Humans and African Great Apes</title><author><name sortKey="Milella, Marco" sort="Milella, Marco" uniqKey="Milella M" first="Marco" last="Milella">Marco Milella</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author></analytic><series><title level="j">PLoS ONE</title><idno type="eISSN">1932-6203</idno><imprint><date when="2014">2014</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Entheseal changes have been widely studied with regard to their correlation to biomechanical stress and their usefulness for biocultural reconstructions. However, anthropological and medical studies have demonstrated the marked influence of both age and sex on the development of these features. Studies of entheseal changes are mostly aimed in testing functional hypotheses and are mostly focused on modern humans, with few data available for non-human primates. The lack of comparative studies on the effect of age and sex on entheseal changes represent a gap in our understanding of the evolutionary basis of both development and degeneration of the human musculoskeletal system. The aim of the present work is to compare age trajectories and patterns of sexual dimorphism in entheseal changes between modern humans and African great apes. To this end we analyzed 23 postcranial entheses in a human contemporary identified skeletal collection (N = 484) and compared the results with those obtained from the analysis of <italic>Pan</italic> (N = 50) and <italic>Gorilla</italic> (N = 47) skeletal specimens. Results highlight taxon-specific age trajectories possibly linked to differences in life history schedules and phyletic relationships. Robusticity trajectories separate <italic>Pan</italic> and modern humans from <italic>Gorilla</italic>, whereas enthesopathic patterns are unique in modern humans and possibly linked to their extended potential lifespan. Comparisons between sexes evidence a decreasing dimorphism in robusticity from <italic>Gorilla</italic>, to modern humans to <italic>Pan</italic>, which is likely linked to the role played by size, lifespan and physical activity on robusticity development. The present study confirms previous hypotheses on the possible relevance of EC in the study of life history, pointing moreover to their usefulness in evolutionary studies.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Lieverse, Ar" uniqKey="Lieverse A">AR Lieverse</name></author><author><name sortKey="Bazaliiskii, Vi" uniqKey="Bazaliiskii V">VI Bazaliiskii</name></author><author><name sortKey="Goriunova, Oi" uniqKey="Goriunova O">OI Goriunova</name></author><author><name sortKey="Weber, Aw" uniqKey="Weber A">AW Weber</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Lieverse, Ar" uniqKey="Lieverse A">AR Lieverse</name></author><author><name sortKey="Bazaliiskii, Vi" uniqKey="Bazaliiskii V">VI Bazaliiskii</name></author><author><name sortKey="Goriunova, Oi" uniqKey="Goriunova O">OI Goriunova</name></author><author><name sortKey="Weber, Aw" uniqKey="Weber A">AW Weber</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Dutour, O" uniqKey="Dutour O">O Dutour</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Molnar, P" uniqKey="Molnar P">P Molnar</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Eshed, V" uniqKey="Eshed V">V Eshed</name></author><author><name sortKey="Gopher, A" uniqKey="Gopher A">A Gopher</name></author><author><name sortKey="Galili, E" uniqKey="Galili E">E Galili</name></author><author><name sortKey="Hershkovitz, I" uniqKey="Hershkovitz I">I Hershkovitz</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Molleson, T" uniqKey="Molleson T">T Molleson</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Lovell, Nc" uniqKey="Lovell N">NC Lovell</name></author><author><name sortKey="Dublenko, Aa" uniqKey="Dublenko A">AA Dublenko</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Henderson, C" uniqKey="Henderson C">C Henderson</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Villotte, S" uniqKey="Villotte S">S Villotte</name></author><author><name sortKey="Knusel, Cj" uniqKey="Knusel C">CJ Knusel</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hawkey, De" uniqKey="Hawkey D">DE Hawkey</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Havelkova, P" uniqKey="Havelkova P">P Havelkova</name></author><author><name sortKey="Hladik, M" uniqKey="Hladik M">M Hladik</name></author><author><name sortKey="Veleminsky, P" uniqKey="Veleminsky P">P Veleminsky</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Havelkova, P" uniqKey="Havelkova P">P Havelkova</name></author><author><name sortKey="Villotte, S" uniqKey="Villotte S">S Villotte</name></author><author><name sortKey="Veleminsky, P" uniqKey="Veleminsky P">P Veleminsky</name></author><author><name sortKey="Polacek, L" uniqKey="Polacek L">L Polacek</name></author><author><name sortKey="Dobisikova, M" uniqKey="Dobisikova M">M Dobisikova</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Villotte, S" uniqKey="Villotte S">S Villotte</name></author><author><name sortKey="Churchill, Se" uniqKey="Churchill S">SE Churchill</name></author><author><name sortKey="Dutour, Oj" uniqKey="Dutour O">OJ Dutour</name></author><author><name sortKey="Henry Gambier, D" uniqKey="Henry Gambier D">D Henry-Gambier</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hawkey, De" uniqKey="Hawkey D">DE Hawkey</name></author><author><name sortKey="Merbs, Cf" uniqKey="Merbs C">CF Merbs</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Steen, Sl" uniqKey="Steen S">SL Steen</name></author><author><name sortKey="Lane, Rw" uniqKey="Lane R">RW Lane</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Mariotti, V" uniqKey="Mariotti V">V Mariotti</name></author><author><name sortKey="Facchini, F" uniqKey="Facchini F">F Facchini</name></author><author><name sortKey="Belcastro, Mg" uniqKey="Belcastro M">MG Belcastro</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Mariotti, V" uniqKey="Mariotti V">V Mariotti</name></author><author><name sortKey="Facchini, F" uniqKey="Facchini F">F Facchini</name></author><author><name sortKey="Belcastro, Mg" uniqKey="Belcastro M">MG Belcastro</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Milella, M" uniqKey="Milella M">M Milella</name></author><author><name sortKey="Belcastro, Mg" uniqKey="Belcastro M">MG Belcastro</name></author><author><name sortKey="Zollikofer, Cp" uniqKey="Zollikofer C">CP Zollikofer</name></author><author><name sortKey="Mariotti, V" uniqKey="Mariotti V">V Mariotti</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Alves Cardoso, F" uniqKey="Alves Cardoso F">F Alves Cardoso</name></author><author><name sortKey="Henderson, Cy" uniqKey="Henderson C">CY Henderson</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Villotte, S" uniqKey="Villotte S">S Villotte</name></author><author><name sortKey="Castex, D" uniqKey="Castex D">D Castex</name></author><author><name sortKey="Couallier, V" uniqKey="Couallier V">V Couallier</name></author><author><name sortKey="Dutour, O" uniqKey="Dutour O">O Dutour</name></author><author><name sortKey="Knusel, Cj" uniqKey="Knusel C">CJ Knüsel</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Niinim Ki, S" uniqKey="Niinim Ki S">S Niinimäki</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Benjamin, M" uniqKey="Benjamin M">M Benjamin</name></author><author><name sortKey="Toumi, H" uniqKey="Toumi H">H Toumi</name></author><author><name sortKey="Suzuki, D" uniqKey="Suzuki D">D Suzuki</name></author><author><name sortKey="Hayashi, K" uniqKey="Hayashi K">K Hayashi</name></author><author><name sortKey="Mcgonagle, D" uniqKey="Mcgonagle D">D McGonagle</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Weiss, E" uniqKey="Weiss E">E Weiss</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Weiss, E" uniqKey="Weiss E">E Weiss</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Villotte, S" uniqKey="Villotte S">S Villotte</name></author><author><name sortKey="Knusel, Cj" uniqKey="Knusel C">CJ Knüsel</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Al Oumaoui, I" uniqKey="Al Oumaoui I">I al-Oumaoui</name></author><author><name sortKey="Jimenez Brobeil, S" uniqKey="Jimenez Brobeil S">S Jimenez-Brobeil</name></author><author><name sortKey="Du Souich, P" uniqKey="Du Souich P">P du Souich</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Wilczak, Ca" uniqKey="Wilczak C">CA Wilczak</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Benjamin, M" uniqKey="Benjamin M">M Benjamin</name></author><author><name sortKey="Kumai, T" uniqKey="Kumai T">T Kumai</name></author><author><name sortKey="Milz, S" uniqKey="Milz S">S Milz</name></author><author><name sortKey="Boszczyk, Bm" uniqKey="Boszczyk B">BM Boszczyk</name></author><author><name sortKey="Boszczyk, Aa" uniqKey="Boszczyk A">AA Boszczyk</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Benjamin, M" uniqKey="Benjamin M">M Benjamin</name></author><author><name sortKey="Ralphs, Jr" uniqKey="Ralphs J">JR Ralphs</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Henderson, Cy" uniqKey="Henderson C">CY Henderson</name></author><author><name sortKey="Mariotti, V" uniqKey="Mariotti V">V Mariotti</name></author><author><name sortKey="Pany Kucera, D" uniqKey="Pany Kucera D">D Pany-Kucera</name></author><author><name sortKey="Villotte, S" uniqKey="Villotte S">S Villotte</name></author><author><name sortKey="Wilczak, C" uniqKey="Wilczak C">C Wilczak</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Robb, Je" uniqKey="Robb J">JE Robb</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Milella, M" uniqKey="Milella M">M Milella</name></author><author><name sortKey="Zollikofer, Cp" uniqKey="Zollikofer C">CP Zollikofer</name></author><author><name sortKey="De Le N, Mp" uniqKey="De Le N M">MP De León</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Morimoto, N" uniqKey="Morimoto N">N Morimoto</name></author><author><name sortKey="Zollikofer, Cp" uniqKey="Zollikofer C">CP Zollikofer</name></author><author><name sortKey="Ponce De Leon, Ms" uniqKey="Ponce De Leon M">MS Ponce de Leon</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Morimoto, N" uniqKey="Morimoto N">N Morimoto</name></author><author><name sortKey="Ponce De Leon, Ms" uniqKey="Ponce De Leon M">MS Ponce de Leon</name></author><author><name sortKey="Zollikofer, Cp" uniqKey="Zollikofer C">CP Zollikofer</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Maggiano, Is" uniqKey="Maggiano I">IS Maggiano</name></author><author><name sortKey="Schultz, M" uniqKey="Schultz M">M Schultz</name></author><author><name sortKey="Kierdorf, H" uniqKey="Kierdorf H">H Kierdorf</name></author><author><name sortKey="Sosa, Ts" uniqKey="Sosa T">TS Sosa</name></author><author><name sortKey="Maggiano, Cm" uniqKey="Maggiano C">CM Maggiano</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Marchi, D" uniqKey="Marchi D">D Marchi</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Marchi, D" uniqKey="Marchi D">D Marchi</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Patel, Ba" uniqKey="Patel B">BA Patel</name></author><author><name sortKey="Ruff, Cb" uniqKey="Ruff C">CB Ruff</name></author><author><name sortKey="Simons, El" uniqKey="Simons E">EL Simons</name></author><author><name sortKey="Organ, Jm" uniqKey="Organ J">JM Organ</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Ruff, Cb" uniqKey="Ruff C">CB Ruff</name></author><author><name sortKey="Runestad, Ja" uniqKey="Runestad J">JA Runestad</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Shaw, Cn" uniqKey="Shaw C">CN Shaw</name></author><author><name sortKey="Stock, Jt" uniqKey="Stock J">JT Stock</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Barak, Mm" uniqKey="Barak M">MM Barak</name></author><author><name sortKey="Lieberman, De" uniqKey="Lieberman D">DE Lieberman</name></author><author><name sortKey="Raichlen, D" uniqKey="Raichlen D">D Raichlen</name></author><author><name sortKey="Pontzer, H" uniqKey="Pontzer H">H Pontzer</name></author><author><name sortKey="Warrener, Ag" uniqKey="Warrener A">AG Warrener</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Ryan, Tm" uniqKey="Ryan T">TM Ryan</name></author><author><name sortKey="Shaw, Cn" uniqKey="Shaw C">CN Shaw</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Shaw, Cn" uniqKey="Shaw C">CN Shaw</name></author><author><name sortKey="Ryan, Tm" uniqKey="Ryan T">TM Ryan</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Tsegai, Zj" uniqKey="Tsegai Z">ZJ Tsegai</name></author><author><name sortKey="Kivell, Tl" uniqKey="Kivell T">TL Kivell</name></author><author><name sortKey="Gross, T" uniqKey="Gross T">T Gross</name></author><author><name sortKey="Nguyen, Nh" uniqKey="Nguyen N">NH Nguyen</name></author><author><name sortKey="Pahr, Dh" uniqKey="Pahr D">DH Pahr</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Desilva, Jm" uniqKey="Desilva J">JM DeSilva</name></author><author><name sortKey="Devlin, Mj" uniqKey="Devlin M">MJ Devlin</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Morimoto, N" uniqKey="Morimoto N">N Morimoto</name></author><author><name sortKey="De Leon, Ms" uniqKey="De Leon M">MS De Leon</name></author><author><name sortKey="Nishimura, T" uniqKey="Nishimura T">T Nishimura</name></author><author><name sortKey="Zollikofer, Cp" uniqKey="Zollikofer C">CP Zollikofer</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Kikuchi, Y" uniqKey="Kikuchi Y">Y Kikuchi</name></author><author><name sortKey="Takemoto, H" uniqKey="Takemoto H">H Takemoto</name></author><author><name sortKey="Kuraoka, A" uniqKey="Kuraoka A">A Kuraoka</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Myatt, Jp" uniqKey="Myatt J">JP Myatt</name></author><author><name sortKey="Crompton, Rh" uniqKey="Crompton R">RH Crompton</name></author><author><name sortKey="Payne Davis, Rc" uniqKey="Payne Davis R">RC Payne-Davis</name></author><author><name sortKey="Vereecke, Ee" uniqKey="Vereecke E">EE Vereecke</name></author><author><name sortKey="Isler, K" uniqKey="Isler K">K Isler</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Jurmain, R" uniqKey="Jurmain R">R Jurmain</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Jurmain, R" uniqKey="Jurmain R">R Jurmain</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Schultz, Ah" uniqKey="Schultz A">AH Schultz</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Schultz, Ah" uniqKey="Schultz A">AH Schultz</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Drapeau, Ms" uniqKey="Drapeau M">MS Drapeau</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Kelley, J" uniqKey="Kelley J">J Kelley</name></author><author><name sortKey="Schwartz, Gt" uniqKey="Schwartz G">GT Schwartz</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Dean, Mc" uniqKey="Dean M">MC Dean</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Schwartz, Gt" uniqKey="Schwartz G">GT Schwartz</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Zollikofer, Cp" uniqKey="Zollikofer C">CP Zollikofer</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Lieberman, De" uniqKey="Lieberman D">DE Lieberman</name></author><author><name sortKey="Carlo, J" uniqKey="Carlo J">J Carlo</name></author><author><name sortKey="Ponce De Le N, M" uniqKey="Ponce De Le N M">M Ponce de León</name></author><author><name sortKey="Zollikofer, Cpe" uniqKey="Zollikofer C">CPE Zollikofer</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Morimoto, N" uniqKey="Morimoto N">N Morimoto</name></author><author><name sortKey="Ogihara, N" uniqKey="Ogihara N">N Ogihara</name></author><author><name sortKey="Katayama, K" uniqKey="Katayama K">K Katayama</name></author><author><name sortKey="Shiota, K" uniqKey="Shiota K">K Shiota</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Bolter, Dr" uniqKey="Bolter D">DR Bolter</name></author><author><name sortKey="Zihlman, Al" uniqKey="Zihlman A">AL Zihlman</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Watts, E" uniqKey="Watts E">E Watts</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Schultz, Ah" uniqKey="Schultz A">AH Schultz</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Zihlman, Al" uniqKey="Zihlman A">AL Zihlman</name></author><author><name sortKey="Bolter, Dr" uniqKey="Bolter D">DR Bolter</name></author><author><name sortKey="Boesch, C" uniqKey="Boesch C">C Boesch</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Ruff, Cb" uniqKey="Ruff C">CB Ruff</name></author><author><name sortKey="Burgess, Ml" uniqKey="Burgess M">ML Burgess</name></author><author><name sortKey="Bromage, Tg" uniqKey="Bromage T">TG Bromage</name></author><author><name sortKey="Mudakikwa, A" uniqKey="Mudakikwa A">A Mudakikwa</name></author><author><name sortKey="Mcfarlin, Sc" uniqKey="Mcfarlin S">SC McFarlin</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Derousseau, Cj" uniqKey="Derousseau C">CJ DeRousseau</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Derousseau, Cj" uniqKey="Derousseau C">CJ DeRousseau</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Derousseau, Cj" uniqKey="Derousseau C">CJ DeRousseau</name></author><author><name sortKey="Rawlins, Rg" uniqKey="Rawlins R">RG Rawlins</name></author><author><name sortKey="Denlinger, Jl" uniqKey="Denlinger J">JL Denlinger</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Pope, Ns" uniqKey="Pope N">NS Pope</name></author><author><name sortKey="Gould, Kg" uniqKey="Gould K">KG Gould</name></author><author><name sortKey="Anderson, Dc" uniqKey="Anderson D">DC Anderson</name></author><author><name sortKey="Mann, Dr" uniqKey="Mann D">DR Mann</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Sumner, Dr" uniqKey="Sumner D">DR Sumner</name></author><author><name sortKey="Morbeck, Me" uniqKey="Morbeck M">ME Morbeck</name></author><author><name sortKey="Lobick, Jj" uniqKey="Lobick J">JJ Lobick</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Morbeck, Me" uniqKey="Morbeck M">ME Morbeck</name></author><author><name sortKey="Galloway, A" uniqKey="Galloway A">A Galloway</name></author><author><name sortKey="Sumner, Dr" uniqKey="Sumner D">DR Sumner</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Schultz, Ah" uniqKey="Schultz A">AH Schultz</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Robson, Sl" uniqKey="Robson S">SL Robson</name></author><author><name sortKey="Wood, B" uniqKey="Wood B">B Wood</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Karasik, D" uniqKey="Karasik D">D Karasik</name></author><author><name sortKey="Kiel, Dp" uniqKey="Kiel D">DP Kiel</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Bogin, Ba" uniqKey="Bogin B">BA Bogin</name></author><author><name sortKey="Smith, Bh" uniqKey="Smith B">BH Smith</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Zollikofer, Cpe" uniqKey="Zollikofer C">CPE Zollikofer</name></author><author><name sortKey="Ponce De Le N, Ms" uniqKey="Ponce De Le N M">MS Ponce de León</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Kaplan, H" uniqKey="Kaplan H">H Kaplan</name></author><author><name sortKey="Hill, K" uniqKey="Hill K">K Hill</name></author><author><name sortKey="Lancaster, J" uniqKey="Lancaster J">J Lancaster</name></author><author><name sortKey="Hurtado, Am" uniqKey="Hurtado A">AM Hurtado</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Leigh, Sr" uniqKey="Leigh S">SR Leigh</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Smith, Rj" uniqKey="Smith R">RJ Smith</name></author><author><name sortKey="Jungers, Wl" uniqKey="Jungers W">WL Jungers</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Wood, B" uniqKey="Wood B">B Wood</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Frayer, Dw" uniqKey="Frayer D">DW Frayer</name></author><author><name sortKey="Wolpoff, Mh" uniqKey="Wolpoff M">MH Wolpoff</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Martin Dupont, S" uniqKey="Martin Dupont S">S Martin-Dupont</name></author><author><name sortKey="Cunha, E" uniqKey="Cunha E">E Cunha</name></author><author><name sortKey="Rouge, D" uniqKey="Rouge D">D Rouge</name></author><author><name sortKey="Crubezy, E" uniqKey="Crubezy E">E Crubezy</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Lovell, Nc" uniqKey="Lovell N">NC Lovell</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Bolter, Dr" uniqKey="Bolter D">DR Bolter</name></author><author><name sortKey="Zihlman, Al" uniqKey="Zihlman A">AL Zihlman</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Molnar, S" uniqKey="Molnar S">S Molnar</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Niinim Ki, S" uniqKey="Niinim Ki S">S Niinimäki</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Leigh, Sr" uniqKey="Leigh S">SR Leigh</name></author><author><name sortKey="Shea, Bt" uniqKey="Shea B">BT Shea</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Doran, Dm" uniqKey="Doran D">DM Doran</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Rothschild, Bm" uniqKey="Rothschild B">BM Rothschild</name></author><author><name sortKey="Woods, Rj" uniqKey="Woods R">RJ Woods</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Jiang, Y" uniqKey="Jiang Y">Y Jiang</name></author><author><name sortKey="Zhao, J" uniqKey="Zhao J">J Zhao</name></author><author><name sortKey="Van Holsbeeck, Mt" uniqKey="Van Holsbeeck M">MT van Holsbeeck</name></author><author><name sortKey="Flynn, Mj" uniqKey="Flynn M">MJ Flynn</name></author><author><name sortKey="Ouyang, X" uniqKey="Ouyang X">X Ouyang</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Milz, S" uniqKey="Milz S">S Milz</name></author><author><name sortKey="Tischer, T" uniqKey="Tischer T">T Tischer</name></author><author><name sortKey="Buettner, A" uniqKey="Buettner A">A Buettner</name></author><author><name sortKey="Schieker, M" uniqKey="Schieker M">M Schieker</name></author><author><name sortKey="Maier, M" uniqKey="Maier M">M Maier</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hoyte, Dan" uniqKey="Hoyte D">DAN Hoyte</name></author><author><name sortKey="Enlow, Dh" uniqKey="Enlow D">DH Enlow</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Dorfl, J" uniqKey="Dorfl J">J Dörfl</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Dorfl, J" uniqKey="Dorfl J">J Dörfl</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Benjamin, M" uniqKey="Benjamin M">M Benjamin</name></author><author><name sortKey="Toumi, H" uniqKey="Toumi H">H Toumi</name></author><author><name sortKey="Suzuki, D" uniqKey="Suzuki D">D Suzuki</name></author><author><name sortKey="Redman, S" uniqKey="Redman S">S Redman</name></author><author><name sortKey="Emery, P" uniqKey="Emery P">P Emery</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Benjamin, M" uniqKey="Benjamin M">M Benjamin</name></author><author><name sortKey="Milz, S" uniqKey="Milz S">S Milz</name></author><author><name sortKey="Bydder, Gm" uniqKey="Bydder G">GM Bydder</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Richmond, Bg" uniqKey="Richmond B">BG Richmond</name></author><author><name sortKey="Jungers, Wl" uniqKey="Jungers W">WL Jungers</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Niinim Ki, S" uniqKey="Niinim Ki S">S Niinimäki</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Peterson, J" uniqKey="Peterson J">J Peterson</name></author></analytic></biblStruct></listBibl></div1></back></TEI><pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">PLoS One</journal-id><journal-id journal-id-type="iso-abbrev">PLoS ONE</journal-id><journal-id journal-id-type="publisher-id">plos</journal-id><journal-id journal-id-type="pmc">plosone</journal-id><journal-title-group><journal-title>PLoS ONE</journal-title></journal-title-group><issn pub-type="epub">1932-6203</issn><publisher><publisher-name>Public Library of Science</publisher-name><publisher-loc>San Francisco, USA</publisher-loc></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">25251439</article-id><article-id pub-id-type="pmc">4175998</article-id><article-id pub-id-type="publisher-id">PONE-D-14-12787</article-id><article-id pub-id-type="doi">10.1371/journal.pone.0107963</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group><subj-group subj-group-type="Discipline-v2"><subject>Biology and Life Sciences</subject><subj-group><subject>Anatomy</subject><subj-group><subject>Musculoskeletal System</subject></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v2"><subject>Social Sciences</subject><subj-group><subject>Anthropology</subject></subj-group></subj-group></article-categories><title-group><article-title>The Influence of Life History and Sexual Dimorphism on Entheseal Changes in Modern Humans and African Great Apes</article-title><alt-title alt-title-type="running-head">Entheseal Changes in Modern Humans and African Great Apes</alt-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Milella</surname><given-names>Marco</given-names></name><xref ref-type="aff" rid="aff1"></xref><xref ref-type="corresp" rid="cor1"><sup>*</sup></xref></contrib></contrib-group><aff id="aff1"><addr-line>Anthropological Institute and Museum, University of Zurich, Zurich, Switzerland</addr-line></aff><contrib-group><contrib contrib-type="editor"><name><surname>Bondioli</surname><given-names>Luca</given-names></name><role>Editor</role><xref ref-type="aff" rid="edit1"></xref></contrib></contrib-group><aff id="edit1"><addr-line>Museo Nazionale Preistorico Etnografico ‘L. Pigorini’, Italy</addr-line></aff><author-notes><corresp id="cor1">* E-mail: <email>marco.milella@aim.uzh.ch</email></corresp><fn fn-type="conflict"><p><bold>Competing Interests: </bold>The author has declared that no competing interests exist.</p></fn><fn fn-type="con"><p>Conceived and designed the experiments: MM. Performed the experiments: MM. Analyzed the data: MM. Contributed reagents/materials/analysis tools: MM. Contributed to the writing of the manuscript: MM.</p></fn></author-notes><pub-date pub-type="collection"><year>2014</year></pub-date><pub-date pub-type="epub"><day>24</day><month>9</month><year>2014</year></pub-date><volume>9</volume><issue>9</issue><elocation-id>e107963</elocation-id><history><date date-type="received"><day>28</day><month>3</month><year>2014</year></date><date date-type="accepted"><day>16</day><month>8</month><year>2014</year></date></history><permissions><copyright-year>2014</copyright-year><copyright-holder>Marco Milella</copyright-holder><license><license-p>This is an open-access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution License</ext-link>, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p></license></permissions><abstract><p>Entheseal changes have been widely studied with regard to their correlation to biomechanical stress and their usefulness for biocultural reconstructions. However, anthropological and medical studies have demonstrated the marked influence of both age and sex on the development of these features. Studies of entheseal changes are mostly aimed in testing functional hypotheses and are mostly focused on modern humans, with few data available for non-human primates. The lack of comparative studies on the effect of age and sex on entheseal changes represent a gap in our understanding of the evolutionary basis of both development and degeneration of the human musculoskeletal system. The aim of the present work is to compare age trajectories and patterns of sexual dimorphism in entheseal changes between modern humans and African great apes. To this end we analyzed 23 postcranial entheses in a human contemporary identified skeletal collection (N = 484) and compared the results with those obtained from the analysis of <italic>Pan</italic> (N = 50) and <italic>Gorilla</italic> (N = 47) skeletal specimens. Results highlight taxon-specific age trajectories possibly linked to differences in life history schedules and phyletic relationships. Robusticity trajectories separate <italic>Pan</italic> and modern humans from <italic>Gorilla</italic>, whereas enthesopathic patterns are unique in modern humans and possibly linked to their extended potential lifespan. Comparisons between sexes evidence a decreasing dimorphism in robusticity from <italic>Gorilla</italic>, to modern humans to <italic>Pan</italic>, which is likely linked to the role played by size, lifespan and physical activity on robusticity development. The present study confirms previous hypotheses on the possible relevance of EC in the study of life history, pointing moreover to their usefulness in evolutionary studies.</p></abstract><funding-group><funding-statement>This author has no support or funding to report.</funding-statement></funding-group><counts><page-count count="10"></page-count></counts><custom-meta-group><custom-meta id="data-availability"><meta-name>Data Availability</meta-name><meta-value>The authors confirm that all data underlying the findings are fully available without restriction. All relevant data are within the paper and its Supporting Information files.</meta-value></custom-meta></custom-meta-group></article-meta><notes><title>Data Availability</title><p>The authors confirm that all data underlying the findings are fully available without restriction. All relevant data are within the paper and its Supporting Information files.</p></notes></front><body><sec id="s1"><title>Introduction</title><p>Due to their supposed link with physical activity, morphological changes at the level of the entheses (skeletal attachment sites of muscles, ligaments and joint capsules), or entheseal changes (EC) <xref rid="pone.0107963-Jurmain1" ref-type="bibr">[1]</xref> have been widely adopted as proxies of <italic>in vivo</italic> biomechanical stress in bioarchaeological studies <xref rid="pone.0107963-Lieverse1" ref-type="bibr">[2]</xref>, <xref rid="pone.0107963-Lieverse2" ref-type="bibr">[3]</xref>–<xref rid="pone.0107963-Lovell1" ref-type="bibr">[8]</xref>. EC have indeed been used to test biocultural hypotheses about differences between contrasting socio-economic systems, possible presence of sexual subdivision of labour in past societies <xref rid="pone.0107963-Henderson1" ref-type="bibr">[9]</xref>–<xref rid="pone.0107963-Villotte2" ref-type="bibr">[14]</xref>, and performance of specific daily physical activities <xref rid="pone.0107963-Lovell1" ref-type="bibr">[8]</xref>,<xref rid="pone.0107963-Hawkey2" ref-type="bibr">[15]</xref>,<xref rid="pone.0107963-Steen1" ref-type="bibr">[16]</xref>. Nonetheless, recent analyses of identified skeletal collections highlighted the multifactorial etiology of EC <xref rid="pone.0107963-Mariotti1" ref-type="bibr">[17]</xref>–<xref rid="pone.0107963-Niinimki1" ref-type="bibr">[23]</xref>. Sexual dimorphism <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>,<xref rid="pone.0107963-Villotte4" ref-type="bibr">[22]</xref>, anatomy of the attachment sites <xref rid="pone.0107963-Villotte3" ref-type="bibr">[21]</xref>,<xref rid="pone.0107963-Villotte4" ref-type="bibr">[22]</xref>, and, especially, age <xref rid="pone.0107963-Mariotti1" ref-type="bibr">[17]</xref>–<xref rid="pone.0107963-Benjamin1" ref-type="bibr">[24]</xref> appear strongly correlated with the expression of EC. Differences in the expression of EC in males and females has been observed in both archaeological and contemporary skeletal samples <xref rid="pone.0107963-Villotte2" ref-type="bibr">[14]</xref>,<xref rid="pone.0107963-Steen1" ref-type="bibr">[16]</xref>,<xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>,<xref rid="pone.0107963-Villotte4" ref-type="bibr">[22]</xref>,<xref rid="pone.0107963-Niinimki1" ref-type="bibr">[23]</xref>,<xref rid="pone.0107963-Weiss1" ref-type="bibr">[25]</xref>–<xref rid="pone.0107963-Wilczak1" ref-type="bibr">[29]</xref>, a pattern attributed to different activities performed by sexes and/or to the effect of physiological factors (e.g. hormones) on the expression of EC. About the relationship between the type of tissue of the attachment sites and EC, medical and anthropological studies pointed to substantial differences in terms of morphological variability between fibrous and fibrocartilaginous entheses (characterized, respectively, by the presence of dense fibrous connective tissue and fibrocartilage at the interface between bone and tendon <xref rid="pone.0107963-Benjamin2" ref-type="bibr">[30]</xref>,<xref rid="pone.0107963-Benjamin3" ref-type="bibr">[31]</xref>) <xref rid="pone.0107963-Villotte3" ref-type="bibr">[21]</xref>,<xref rid="pone.0107963-Benjamin2" ref-type="bibr">[30]</xref>,<xref rid="pone.0107963-Villotte6" ref-type="bibr">[32]</xref>. Such distinction has been recently adopted in anthropology, and considerations on the anatomy of entheses have been fully integrated in both the stages of data collection and discussion of results <xref rid="pone.0107963-AlvesCardoso1" ref-type="bibr">[20]</xref>,<xref rid="pone.0107963-Villotte4" ref-type="bibr">[22]</xref>,<xref rid="pone.0107963-Villotte6" ref-type="bibr">[32]</xref>,<xref rid="pone.0107963-Henderson2" ref-type="bibr">[33]</xref>. Concerning age, both medical and anthropological studies agree on the strong role played by this variable on the expression of EC <xref rid="pone.0107963-Mariotti1" ref-type="bibr">[17]</xref>,<xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>–<xref rid="pone.0107963-Villotte3" ref-type="bibr">[21]</xref>,<xref rid="pone.0107963-Niinimki1" ref-type="bibr">[23]</xref>,<xref rid="pone.0107963-Benjamin1" ref-type="bibr">[24]</xref>,<xref rid="pone.0107963-Durigon1" ref-type="bibr">[34]</xref>,<xref rid="pone.0107963-Robb1" ref-type="bibr">[35]</xref>, a complex phenomenon likely related to several causes, including the response of bone to continuous microtraumatic stress related to daily biomechanical stimuli, ontogenetic processes of the musculoskeletal system, and degenerative processes associated with aging.</p><p>Such results, besides suggesting a revision of previous functional hypotheses, allow placing EC in a wider perspective, establishing a link between these features and biological processes such as the evolution and development of the musculoskeletal system as well as shared and derived patterns in the life history of primates <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>,<xref rid="pone.0107963-Milella2" ref-type="bibr">[36]</xref>.</p><p>Geometric properties of long bones <xref rid="pone.0107963-Morimoto1" ref-type="bibr">[37]</xref>–<xref rid="pone.0107963-Shaw1" ref-type="bibr">[44]</xref>, trabecular architecture of various skeletal regions <xref rid="pone.0107963-Barak1" ref-type="bibr">[45]</xref>–<xref rid="pone.0107963-DeSilva1" ref-type="bibr">[49]</xref>, organization of postcranial muscles <xref rid="pone.0107963-Morimoto3" ref-type="bibr">[50]</xref>–<xref rid="pone.0107963-Myatt1" ref-type="bibr">[52]</xref> and patterns of joint diseases, trauma and various pathological conditions <xref rid="pone.0107963-Jurmain2" ref-type="bibr">[53]</xref>–<xref rid="pone.0107963-Schultz2" ref-type="bibr">[56]</xref> represent some of the main topics of anthropological research on the musculoskeletal system of primates and are mainly framed in a functional perspective. On the other hand, despite the wide array of studies on EC in modern humans, few data are available on EC in non-human primates, with only one publication <xref rid="pone.0107963-Drapeau1" ref-type="bibr">[57]</xref> focusing on interspecific patterns of bilateral asymmetry and forelimb-hindlimb ratios, also in this case investigated in order to test their reliability in reflecting intertaxa behavioral patterns.</p><p>As far as life history is concerned, observations on the skeleton have been normally focused on early ontogenetic stages through the analysis of dental data <xref rid="pone.0107963-Kelley1" ref-type="bibr">[58]</xref>–<xref rid="pone.0107963-Schwartz1" ref-type="bibr">[60]</xref> as well as cranial <xref rid="pone.0107963-Zollikofer1" ref-type="bibr">[61]</xref>–<xref rid="pone.0107963-Bolter1" ref-type="bibr">[64]</xref> and postcranial <xref rid="pone.0107963-Bolter1" ref-type="bibr">[64]</xref>–<xref rid="pone.0107963-Ruff2" ref-type="bibr">[68]</xref> developmental schedules. Topics of such studies include analyses of interspecific comparisons of skeletal growth patterns <xref rid="pone.0107963-Bolter1" ref-type="bibr">[64]</xref>, differences in skeletal growth between captive and wild specimens <xref rid="pone.0107963-Zihlman1" ref-type="bibr">[67]</xref>, timing of first permanent molar emergence between wild great apes <xref rid="pone.0107963-Kelley1" ref-type="bibr">[58]</xref> and patterns of evolutionary developmental reorganization of the hominin skull <xref rid="pone.0107963-Zollikofer1" ref-type="bibr">[61]</xref>. Literature on skeletal changes associated to life history variables in the adult is, on the other hand, quite scanty, being represented by few studies mostly focused on aging patterns in single taxa or isolated skeletal samples <xref rid="pone.0107963-DeRousseau1" ref-type="bibr">[69]</xref>–<xref rid="pone.0107963-Schultz4" ref-type="bibr">[75]</xref>. Life history variables in the adult such as gestation length, interbirth interval, post-reproductive survivorship and differential longevity have been mostly studied on living specimens <xref rid="pone.0107963-vanSchaik1" ref-type="bibr">[76]</xref>,<xref rid="pone.0107963-Robson1" ref-type="bibr">[77]</xref>, and rarely used in the discussion of skeletal data. However, clinical research has demonstrated the mutual physiological continuity between the muscular and skeletal systems and their sensibility to both endocrinal and metabolic factors <xref rid="pone.0107963-Karasik1" ref-type="bibr">[78]</xref>. Accordingly, it is expected that entheses, representing the interface between those systems, can be a good candidate for the study of skeletal changes related to life history variables in the adult. This was preliminary confirmed by the recent analysis of a large identified skeletal collection, which highlighted complex age- and sex-specific patterns in EC <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>. Starting from these premises, the aim of the present work is to contribute to the actual knowledge on EC across hominid taxa, by explicitly avoiding a functional perspective, and to explore their usefulness in the study of life history of primates and, more generally, in the analyses of evolutionary processes underlying both growth and aging of the musculosketal system. To this aim, we analyzed patterns of EC in a skeletal sample of modern humans and African great apes, testing the following hypotheses:</p><list list-type="order"><list-item><p>Due to interspecific differences in developmental schedules, interbirth interval and mean potential lifespan <xref rid="pone.0107963-Zollikofer1" ref-type="bibr">[61]</xref>,<xref rid="pone.0107963-Robson1" ref-type="bibr">[77]</xref>,<xref rid="pone.0107963-Bogin1" ref-type="bibr">[79]</xref>–<xref rid="pone.0107963-Robson2" ref-type="bibr">[82]</xref>, we expect modern humans to exhibit diverging age trajectories of EC when compared with African great apes. Specifically, we expect a delayed development of entheseal structures in modern humans due to their delayed somatic growth when compared with African great apes <xref rid="pone.0107963-Zollikofer2" ref-type="bibr">[81]</xref>,<xref rid="pone.0107963-Kaplan1" ref-type="bibr">[83]</xref>,<xref rid="pone.0107963-Leigh1" ref-type="bibr">[84]</xref>.</p></list-item><list-item><p>Due to different levels of sexual dimorphism with regard to body mass and body size <xref rid="pone.0107963-Smith1" ref-type="bibr">[85]</xref>–<xref rid="pone.0107963-Frayer1" ref-type="bibr">[87]</xref>, we expect higher differences between sexes in entheseal development in <italic>Gorilla</italic> when compared with both modern humans and <italic>Pan</italic>.</p></list-item></list></sec><sec sec-type="materials|methods" id="s2"><title>Materials and Methods</title><p>Our sample includes 484 modern humans, 50 <italic>Pan</italic>, and 47 <italic>Gorilla</italic> (<xref ref-type="table" rid="pone-0107963-t001">Table 1</xref>). All specimens were selected on the basis of the absence of pathologies possibly affecting EC (Diffuse Idiopathic Skeletal Hyperostosis - DISH, and spondyloarthropathies) or the normal biomechanics of the body (fractures, dislocations, and dysplasias). DISH and spondyloarthropathies were diagnosed according Rogers & Waldron <xref rid="pone.0107963-Rogers1" ref-type="bibr">[88]</xref> and Martin-Dupont et al. <xref rid="pone.0107963-MartinDupont1" ref-type="bibr">[89]</xref>. Fractures, dislocations and dysplasias were diagnosed according to Lovell <xref rid="pone.0107963-Lovell2" ref-type="bibr">[90]</xref> and Ortner & Putschar <xref rid="pone.0107963-Ortner1" ref-type="bibr">[91]</xref>. The modern humans sample was selected from the identified skeletal collection Frassetto of Sassari <xref rid="pone.0107963-Facchini1" ref-type="bibr">[92]</xref>, housed at the Museum of anthropology of the University of Bologna (Italy). It includes 274 male and 210 female subjects with documented age at death, sex, and profession from the beginning of the 20th century. Only subjects with age at death equal to or greater than twenty years old <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref> were used in this study. Data on African great apes were collected at the Anthropological Institute of the University of Zurich (Switzerland) and the Royal Central African Museum of Tervuren (Belgium). The curators of the collections allowed access to the specimens and use of the relative datasets. Numbers of specimens, together with details on their age and sex can be found in <xref ref-type="supplementary-material" rid="pone.0107963.s014">Appendices S1</xref>–<xref ref-type="supplementary-material" rid="pone.0107963.s015">S2</xref>. The Zurich dataset includes 26 <italic>Gorilla</italic> (N males = 14, N females = 12) and 24 <italic>Pan</italic> (N males = 7, N females = 17), while the Tervuren dataset includes 21 <italic>Gorilla</italic> (N males = 9, N females = 12) and 26 <italic>Pan</italic> (N males = 15, N females = 11). The resulting sample includes both juvenile (erupted M2) and adult (erupted M3) individuals of both sexes (<xref ref-type="table" rid="pone-0107963-t001">Table 1</xref>). While the <italic>Gorilla</italic> dataset includes mostly wild-shot specimens (only two captive individuals), 16 out of 50 <italic>Pan</italic> skeletal specimens are of captive individuals, mostly distributed in the first four age classes (<xref ref-type="supplementary-material" rid="pone.0107963.s015">Appendix S2</xref>). The choice to include in our dataset also captive animals was primarily dictated by the need to maximize the African great apes sample size. However, considering the different life history characterizing wild vs. captive individuals <xref rid="pone.0107963-Watts1" ref-type="bibr">[65]</xref>,<xref rid="pone.0107963-Bolter2" ref-type="bibr">[93]</xref>, we decided to control for such possible source of bias in our analyses (see <xref ref-type="sec" rid="s2">Methods</xref> section).</p><table-wrap id="pone-0107963-t001" orientation="portrait" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0107963.t001</object-id><label>Table 1</label><caption><title>Sample size, sex and age distribution of <italic>Gorilla</italic>, <italic>Pan</italic>, and <italic>Homo</italic>.</title></caption><alternatives><graphic id="pone-0107963-t001-1" xlink:href="pone.0107963.t001"></graphic><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col></colgroup><thead><tr><td align="left" rowspan="1" colspan="1"></td><td colspan="2" align="left" rowspan="1"><italic>Gorilla</italic></td><td colspan="2" align="left" rowspan="1"><italic>Pan</italic></td><td colspan="2" align="left" rowspan="1"><italic>Homo</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">Age class</td><td align="left" rowspan="1" colspan="1">F</td><td align="left" rowspan="1" colspan="1">M</td><td align="left" rowspan="1" colspan="1">F</td><td align="left" rowspan="1" colspan="1">M</td><td align="left" rowspan="1" colspan="1">F</td><td align="left" rowspan="1" colspan="1">M</td></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">8</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">9</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">0</td></tr><tr><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">61</td><td align="left" rowspan="1" colspan="1">64</td></tr><tr><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">4</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">7</td><td align="left" rowspan="1" colspan="1">4</td><td align="left" rowspan="1" colspan="1">42</td><td align="left" rowspan="1" colspan="1">39</td></tr><tr><td align="left" rowspan="1" colspan="1">3</td><td align="left" rowspan="1" colspan="1">8</td><td align="left" rowspan="1" colspan="1">8</td><td align="left" rowspan="1" colspan="1">3</td><td align="left" rowspan="1" colspan="1">9</td><td align="left" rowspan="1" colspan="1">37</td><td align="left" rowspan="1" colspan="1">56</td></tr><tr><td align="left" rowspan="1" colspan="1">4</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">24</td><td align="left" rowspan="1" colspan="1">50</td></tr><tr><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">6</td><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">11</td><td align="left" rowspan="1" colspan="1">32</td></tr><tr><td align="left" rowspan="1" colspan="1">6</td><td align="left" rowspan="1" colspan="1">3</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">22</td><td align="left" rowspan="1" colspan="1">22</td></tr><tr><td align="left" rowspan="1" colspan="1">7</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">13</td><td align="left" rowspan="1" colspan="1">11</td></tr><tr><td align="left" rowspan="1" colspan="1">Total</td><td align="left" rowspan="1" colspan="1">24</td><td align="left" rowspan="1" colspan="1">23</td><td align="left" rowspan="1" colspan="1">28</td><td align="left" rowspan="1" colspan="1">22</td><td align="left" rowspan="1" colspan="1">210</td><td align="left" rowspan="1" colspan="1">274</td></tr></tbody></table></alternatives><table-wrap-foot><fn id="nt101"><label></label><p>F = females; M = males.</p></fn></table-wrap-foot></table-wrap><p>The modern humans sample was separated into classes of ten years each, for a total of seven age classes, while for African great apes we used seven stages of tooth wear (employing the first seven stages of Molnar <xref rid="pone.0107963-Molnar2" ref-type="bibr">[94]</xref> as a proxy for adult age, adding an eighth class – class “0”- for the juveniles, in order to distinguish them from the rest of the sample). We analyzed 23 postcranial entheses (<xref ref-type="table" rid="pone-0107963-t002">Table 2</xref>) with regard to their development in robusticity (surface roughness) as well as proliferative (enthesophytes - EF) and resorptive (osteolytic - OL) entheseal changes (enthesopathies), according to the scoring schemes proposed by Mariotti et al. <xref rid="pone.0107963-Mariotti1" ref-type="bibr">[17]</xref>,<xref rid="pone.0107963-Mariotti2" ref-type="bibr">[18]</xref>. Note that these methods, unlike others <xref rid="pone.0107963-Villotte6" ref-type="bibr">[32]</xref>,<xref rid="pone.0107963-Henderson3" ref-type="bibr">[95]</xref> do not take into account the type of enthesis (i.e. fibrous vs. fibrocartilaginous), a factor often considered in studies on EC <xref rid="pone.0107963-Lieverse1" ref-type="bibr">[2]</xref>,<xref rid="pone.0107963-AlvesCardoso1" ref-type="bibr">[20]</xref>–<xref rid="pone.0107963-Villotte4" ref-type="bibr">[22]</xref>,<xref rid="pone.0107963-Niinimki2" ref-type="bibr">[96]</xref>. Nonetheless, it was chosen for: (a) the previous experience of the author with this method <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>,<xref rid="pone.0107963-Milella2" ref-type="bibr">[36]</xref>,<xref rid="pone.0107963-Milella3" ref-type="bibr">[97]</xref>,<xref rid="pone.0107963-Milella4" ref-type="bibr">[98]</xref>; (b) the site-specificity of the method for scoring robusticity (therefore indirectly taking into account the anatomy of each enthesis), and (c) the chance to consider separately different variables (robusticity and enthesopathies). We considered only the entheseal sites originally included in the method of Mariotti and colleagues <xref rid="pone.0107963-Mariotti1" ref-type="bibr">[17]</xref>. This choice was driven (1) by the scoring criteria for robusticity, which are specific for each site, (2) by the need to analyze different sites in order to check for the expression of EC in the overall postcranial skeleton (3) by the need to compare data collected in different time frames.</p><table-wrap id="pone-0107963-t002" orientation="portrait" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0107963.t002</object-id><label>Table 2</label><caption><title>Entheses considered in the present study: anatomical location and relative functional complex.</title></caption><alternatives><graphic id="pone-0107963-t002-2" xlink:href="pone.0107963.t002"></graphic><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col></colgroup><thead><tr><td align="left" rowspan="1" colspan="1">Enthesis</td><td align="left" rowspan="1" colspan="1">Site</td><td align="left" rowspan="1" colspan="1">Functional complex</td></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1">Costoclavicular lig.</td><td align="left" rowspan="1" colspan="1">clavicle</td><td align="left" rowspan="1" colspan="1">Shoulder</td></tr><tr><td align="left" rowspan="1" colspan="1">Conoid lig.</td><td align="left" rowspan="1" colspan="1">clavicle</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">Trapezoid lig.</td><td align="left" rowspan="1" colspan="1">clavicle</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. pectoralis major</italic></td><td align="left" rowspan="1" colspan="1">clavicle (o)</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. deltoideus</italic></td><td align="left" rowspan="1" colspan="1">clavicle (o)</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. pectoralis major</italic></td><td align="left" rowspan="1" colspan="1">humerus (i)</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. latissimus dorsi/teres major</italic></td><td align="left" rowspan="1" colspan="1">humerus (i)</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. deltoideus</italic></td><td align="left" rowspan="1" colspan="1">humerus (i)</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. triceps brachii</italic></td><td align="left" rowspan="1" colspan="1">scapula (o)</td><td align="left" rowspan="1" colspan="1">Elbow (flexion/extension)</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. brachioradialis</italic></td><td align="left" rowspan="1" colspan="1">humerus (o)</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. biceps brachii</italic></td><td align="left" rowspan="1" colspan="1">radius (i)</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. triceps brachii</italic></td><td align="left" rowspan="1" colspan="1">ulna (i)</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. brachialis</italic></td><td align="left" rowspan="1" colspan="1">ulna (i)</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M pronator teres</italic></td><td align="left" rowspan="1" colspan="1">radius (i)</td><td align="left" rowspan="1" colspan="1">Forearm (pronation/supination)</td></tr><tr><td align="left" rowspan="1" colspan="1">Interosseous membrane</td><td align="left" rowspan="1" colspan="1">radius</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. supinator</italic></td><td align="left" rowspan="1" colspan="1">ulna (o)</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">Enthesis</td><td align="left" rowspan="1" colspan="1">Site</td><td align="left" rowspan="1" colspan="1">Functional complex</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. gluteus maximus</italic></td><td align="left" rowspan="1" colspan="1">femur (i)</td><td align="left" rowspan="1" colspan="1">Hip</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. iliopsoas</italic></td><td align="left" rowspan="1" colspan="1">femur (i)</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. vastus medialis</italic></td><td align="left" rowspan="1" colspan="1">femur (o)</td><td align="left" rowspan="1" colspan="1">Knee</td></tr><tr><td align="left" rowspan="1" colspan="1">Quadriceps tendon</td><td align="left" rowspan="1" colspan="1">tibia</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">Quadriceps tendon</td><td align="left" rowspan="1" colspan="1">patella</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>M. soleus</italic></td><td align="left" rowspan="1" colspan="1">tibia (o)</td><td align="left" rowspan="1" colspan="1">Foot</td></tr><tr><td align="left" rowspan="1" colspan="1">Achilles tendon</td><td align="left" rowspan="1" colspan="1">calcaneus</td><td align="left" rowspan="1" colspan="1"></td></tr></tbody></table></alternatives><table-wrap-foot><fn id="nt102"><label></label><p>M. = muscle; lig. = ligament; o = origin; i = insertion.</p></fn></table-wrap-foot></table-wrap><p>EF and OL were scored according to a four-degree scale <xref rid="pone.0107963-Mariotti1" ref-type="bibr">[17]</xref>–<xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>. In EF, this corresponds to a range from absence of proliferative patterns to the presence of obvious bony spurs. In OL, the scale ranges from absence of resorptive patterns to marked erosive areas at the level of entheses. In analyzing our data, we followed the protocol detailed in Milella et al. <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref> although applying some minor changes. In regression analyses (see below) we used the original four-degree scale for each feature. On the other hand, in pairwise comparison we considered for EF only the presence of obvious proliferative patterns (degrees 2 and 3) while, for OL, the complexity of these features suggested to consider their absence (0), the presence of fine porosities (“pitting” - degree 1) and the presence of extensive osteolytic areas (“erosions” - degrees 2 and 3) <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>. Robusticity was scored according to a five-degree scale (from 0 – extremely weak development– to 4 – marked development). In analysing robusticity, we assumed that each degree represents an underlying continuous variable. Accordingly, in order to explore general patterns of robusticity we grouped the attachment sites into eight functional complexes (<xref ref-type="table" rid="pone-0107963-t002">Table 2</xref>) and calculated a mean robusticity score (MRS) for each complex (see for details Mariotti et al. <xref rid="pone.0107963-Mariotti1" ref-type="bibr">[17]</xref> and Milella et al. <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>, and, for a similar approach, Niinimäki <xref rid="pone.0107963-Niinimki2" ref-type="bibr">[96]</xref>). Qualitative scores like the one used in this study are typically affected by varying degrees of intra- and interobserver error. The method of Mariotti and colleagues for the scoring of EF and OL is reported to be associated with a good (less than 5 percent) intra- and interobserver error for both EF and OL <xref rid="pone.0107963-Mariotti2" ref-type="bibr">[18]</xref>. About robusticity, Mariotti and colleagues report intraobserver and interobserver error percentages of, respectively, 20 and 28 after lumping the first three degrees of robusticity. In this study, we decided to check the intraobserver error for both enthesopathies (EF and OL) and mean robusticity scores (MRS). For the interobserver error, we will discuss the values published by Mariotti and colleagues <xref rid="pone.0107963-Mariotti1" ref-type="bibr">[17]</xref>,<xref rid="pone.0107963-Mariotti2" ref-type="bibr">[18]</xref> on the basis of our results.</p><p>Since a study of bilateral asymmetry is beyond the scope of this work, all analyses were performed only for the left side (chosen in order to maximize the sample size).</p><p>Our analyses included:</p><list list-type="order"><list-item><p>A correlation test between age and EC (EF, OL and MRS) through the Spearman rank sum test.</p></list-item><list-item><p>A test of sexual differences in EC through intra-group pairwise comparison of MRS (Wilcoxon test) and frequencies of EF and OL (chi squared test).</p></list-item></list><p>Given the small size of the African great apes sample, step 1) was computed by pooling together the sexes, while in step 2) we grouped together the age classes 1–7, excluding the juvenile specimens.</p><p>In order to calculate the intraobserver error in the scoring of EF, OL, and MRS, we collected EC data on 58 specimens (20 modern humans, 20 <italic>Pan</italic>, and 18 <italic>Gorilla</italic>) twice, in two independent sessions. The percentage of disagreement between the two sessions and the relative Cohen’s weighted kappa was used for calculating the intraobserver error in the scoring of EF and OL. Intraobserver error in the scoring of MRS was calculated for each complex by comparing the scores from the two sessions with the Wilcoxon test.</p><p>Finally, due to the possible bias introduced on the correlation between EC and age by the presence in the <italic>Pan</italic> sample of a relatively large proportion of captive individuals, we decided to compare the results obtained by analyzing separately captive and wild-shot individuals. A similar comparison was not considered necessary for the <italic>Gorilla</italic> sample, given the small percentage of captive individuals in this group (2/47: 4.3%).</p><p>All statistical analyses were performed with JMP 10.0.0 (SAS Institute Inc. 2012) setting alpha to 95%. Cohen’s weighted kappa was calculated with the package psych (version 1.4.5) <xref rid="pone.0107963-Revelle1" ref-type="bibr">[99]</xref> in R 3.0.3 <xref rid="pone.0107963-R1" ref-type="bibr">[100]</xref>.</p></sec><sec id="s3"><title>Results</title><sec id="s3a"><title>Intraobserver error</title><p>The intraobserver error for EF and OL is acceptable, with error frequencies of, respectively, seven and six percent. The Wilcoxon test performed on the MRS data of the two scoring session reflect overall a negligible disagreement for all the functional complexes (<xref ref-type="supplementary-material" rid="pone.0107963.s001">Table S1</xref>).</p></sec><sec id="s3b"><title>Age</title><p><xref ref-type="supplementary-material" rid="pone.0107963.s002">Tables S2</xref>–<xref ref-type="supplementary-material" rid="pone.0107963.s004">S4</xref> show the summary statistics for EF, OL, and MRS in our sample. Original data for each variable can be found in <xref ref-type="supplementary-material" rid="pone.0107963.s016">Appendices S3</xref>–<xref ref-type="supplementary-material" rid="pone.0107963.s018">S5</xref>.</p><p>In testing interspecific differences in EC age trajectories, we first analyzed only adult individuals, and in a second phase, we included African great apes juvenile specimens.</p><p>In <italic>Pan</italic>, correlation tests between EC and age in captive and wild-shot <italic>Pan</italic> evidence a general lack of differences between these two groups (<xref ref-type="supplementary-material" rid="pone.0107963.s005">Tables S5</xref>–<xref ref-type="supplementary-material" rid="pone.0107963.s007">S7</xref>), justifying their inclusion in a single sample.</p><p>When considering only adult specimens, modern humans are characterized by a positive correlation between EF and age, which is significant for a large number of sites. On the other hand, no significant correlation was found in African great apes between EF and age (<xref ref-type="supplementary-material" rid="pone.0107963.s008">Table S8</xref>; see also <xref ref-type="fig" rid="pone-0107963-g001">Figure 1</xref>). Regarding OL, both positive and negative correlations with age are seen in modern humans (<xref ref-type="supplementary-material" rid="pone.0107963.s009">Table S9</xref>), a pattern that is consistent with earlier analyses <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref> showing a complex pattern of OL, with some sites characterized by different age trajectories of degree 1 and degrees 2 and 3. African great apes are characterized overall by a negative correlation between OL and age. OL correlation with age characterizes a limited suite of sites, only partially overlapping between taxa (<xref ref-type="supplementary-material" rid="pone.0107963.s009">Table S9</xref>). Interestingly, no differences in age trajectories between OL degrees are present in both <italic>Pan</italic> and <italic>Gorilla</italic>. The addition of juvenile specimens does not alter the age patterns in both types of enthesopathic changes (see also <xref ref-type="fig" rid="pone-0107963-g001">Figure 1</xref>).</p><fig id="pone-0107963-g001" orientation="portrait" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0107963.g001</object-id><label>Figure 1</label><caption><title>Box plots of mean EF vs. age.</title><p>Note the clear increase with age of EF in modern humans, and the lack of clear patterns in African great apes.</p></caption><graphic xlink:href="pone.0107963.g001"></graphic></fig><p>As far as robusticity is concerned, results show a significant positive correlation between age and MRS in both <italic>Gorilla</italic> and <italic>Homo</italic>, whereas in <italic>Pan</italic> a similar trend is shown only by complexes of the upper limb and by the foot (<xref ref-type="supplementary-material" rid="pone.0107963.s010">Table S10</xref>). When comparing the hindlimb and forelimb correlation with age, African great apes show on average higher correlation values for the latter, whereas in modern humans no clear difference is evident between the upper and lower limbs.</p><p>MRS age trajectories and rates of MRS increase by age class are specific among taxa (<xref ref-type="fig" rid="pone-0107963-g002">Figure 2</xref>). These include: (a) a fast increase until the fourth age stage in <italic>Gorilla</italic> followed by a sudden decrease; (b) a similar increase in <italic>Pan</italic> until the third age phase, starting from which, both the upper and lower limbs show a less pronounced increase of MRS with advancing age and (c) an increase of MRS in modern humans during the first two age phases (until forty years old) followed by a further growth that reaches a peak around the fifth age phase (between sixty and sixty nine years old) and, starting from the sixth age phase, a slight decrease. When adding the juvenile specimens to the African great apes sample results in a more marked correlation between age and MRS, a pattern that is particularly evident in <italic>Pan</italic> and, overall, represented by steeper age trajectories in both taxa. Comparing such trajectories with those characterizing modern humans, the latter show similar trends, but shifted by about a decade. When excluding from the <italic>Pan</italic> sample the captive specimens, results show a more complex age trajectory of MRS, with an increase of MRS along the first two age phases followed by a stasis and a further increase starting from the fourth age class (<xref ref-type="fig" rid="pone-0107963-g003">Figure 3</xref>).</p><fig id="pone-0107963-g002" orientation="portrait" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0107963.g002</object-id><label>Figure 2</label><caption><title>Box plots of mean robusticity score (MRS) vs. age.</title><p>Note the steep and regular increase of MRS in <italic>Gorilla</italic>, and the more complex trajectories shared by <italic>Pan</italic> and modern humans.</p></caption><graphic xlink:href="pone.0107963.g002"></graphic></fig><fig id="pone-0107963-g003" orientation="portrait" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0107963.g003</object-id><label>Figure 3</label><caption><title><italic>Pan</italic>: mean robusticity score (MRS) vs. age after excluding the captive specimens (grey line and box plots) compared with the age trajectories of the complete <italic>Pan</italic> sample (dotted black line, see also <xref ref-type="fig" rid="pone-0107963-g002"><bold>Figure 2</bold></xref>).</title><p>Note the increase in complexity of the age trajectories characterizing the wild specimens.</p></caption><graphic xlink:href="pone.0107963.g003"></graphic></fig></sec><sec id="s3c"><title>Sexual dimorphism</title><p>Given the observed correlation between age and EC we did a preliminary check for age differences among taxa (only adult specimens were considered in testing for EC sexual dimorphism). Intersex comparison for mean age in modern humans (by a Student t test) and sample size of each age class in <italic>Pan</italic> (by a Chi square test) resulted in no significant differences. In <italic>Gorilla,</italic> the age classes 2, 4, and 5 have different sample sizes between sexes, resulting in males being on average older than females, and, therefore, analyses of sexual dimorphism in this taxon are more prone to be biased by the effect of age. Despite this issue, we preferred to compare sexes without subdividing the sample into age classes for all taxa, and to consider the possible effect of age on <italic>Gorilla</italic> sexual dimorphism when discussing our results. In modern humans significant differences between the sexes in EF are limited to the lower limb, with males showing higher frequencies at the mm. <italic>quadriceps femoris</italic> (patellar site) and <italic>soleus.</italic> In the African great apes sample, a test of differences between the sexes for EF was possible only for the ulnar enthesis of the m. <italic>tricipes brachii</italic> (<italic>Gorilla</italic> and <italic>Pan</italic>) and for the quadriceps tendon on the patella (<italic>Gorilla</italic>), which overall show a lack of differences between sexes (<xref ref-type="supplementary-material" rid="pone.0107963.s011">Table S11</xref>). For OL, these are more present in males for the costoclavicular ligament, the m. <italic>biceps brachii</italic>, and for the m. <italic>pectoralis major</italic> (humeral site). The same pattern is present in <italic>Gorilla</italic> for the costoclavicular ligament, while no sexual differences were found in <italic>Pan</italic> (<xref ref-type="supplementary-material" rid="pone.0107963.s012">Table S12</xref>). Analysis of sexual dimorphism in MRS reveals higher development in modern human males for the elbow, forearm, hip, and for the overall upper limb. In <italic>Gorilla,</italic> males show higher development at the elbow, knee, and for the overall upper and lower limbs, while no sexual dimorphism was observed in <italic>Pan</italic> (<xref ref-type="supplementary-material" rid="pone.0107963.s013">Table S13</xref>).</p></sec></sec><sec id="s4"><title>Discussion</title><sec id="s4a"><title>Intraobserver error</title><p>Intraobserver error for the scoring of EF, OL, and MRS reflect a good consistency among our data. Interobserver errors for EF and OL have been reported to correspond to less than 5% <xref rid="pone.0107963-Mariotti2" ref-type="bibr">[18]</xref>, whereas for robusticity the same error (but considering only three robusticity degrees) has shown to reach 20% <xref rid="pone.0107963-Mariotti1" ref-type="bibr">[17]</xref>. In our case no significant intraobserver disagreement was found for MRS. At the same time a calculation of intraobserver error of robusticity scores for single sites gives a value of 41%, but with a root mean squared error of only 0.9. In fact, most disagreements are due to differences in scoring of just one degree of robusticity, as already noted by Mariotti and colleagues <xref rid="pone.0107963-Mariotti1" ref-type="bibr">[17]</xref>. Such results stress the advantage of using functional mean robusticity scores (MRS) instead of raw data from single sites, and, at the same time, suggest that the interobserver reported in the literature <xref rid="pone.0107963-Mariotti1" ref-type="bibr">[17]</xref> for this variable would be markedly lower when using MRS.</p></sec><sec id="s4b"><title>Age</title><p>Our first hypothesis postulated taxon-specific age trajectories of EC, and, specifically, a delayed increase of EC with age in modern humans when compared with African great apes.</p><p>This hypothesis was partially confirmed by our results, which show: (a) a linear increase of MRS with age in <italic>Gorilla,</italic> (b) a more complex and, starting from the third age phase, more moderate increase in <italic>Pan</italic> and modern humans, and, (c) a delayed increase of MRS growth as a function of absolute age in modern humans. Such results are consistent with interspecific differences in growth rate <xref rid="pone.0107963-Robson1" ref-type="bibr">[77]</xref>,<xref rid="pone.0107963-Zollikofer2" ref-type="bibr">[81]</xref>,<xref rid="pone.0107963-Leigh1" ref-type="bibr">[84]</xref>,<xref rid="pone.0107963-Leigh2" ref-type="bibr">[101]</xref>,<xref rid="pone.0107963-Doran1" ref-type="bibr">[102]</xref>. In <italic>Gorilla</italic>, the high rate of MRS increase in this taxon and its delayed decrease when compared with <italic>Pan</italic> can be traced back to Gorillas’ fast somatic growth <xref rid="pone.0107963-Robson1" ref-type="bibr">[77]</xref> and, at least in males, a later slowing of growth when compared with <italic>Pan</italic><xref rid="pone.0107963-Leigh2" ref-type="bibr">[101]</xref>. Age trajectories of <italic>Pan</italic> and modern humans are more complex. The high rate of MRS increase in <italic>Pan</italic> during early age overlaps that observed in <italic>Gorilla</italic> and is consistent with the faster somatic growth of this taxon compared to modern humans <xref rid="pone.0107963-Robson1" ref-type="bibr">[77]</xref>. In both <italic>Pan</italic> and modern humans the MRS trajectories are similar and characterized by an increase until the second age phase, a stasis until the third, and a further increase. Such results are further strengthened when considering only wild <italic>Pan</italic> specimens. The resulting trajectories are indeed more complex and similar to those of modern humans, with the presence of a marked stasis in MRS increase starting from the second age phase. Such pattern can be linked to the exclusion of captive specimens, which, due to their faster somatic growth <xref rid="pone.0107963-Watts1" ref-type="bibr">[65]</xref>,<xref rid="pone.0107963-Bolter2" ref-type="bibr">[93]</xref>, are likely to bias the MRS patterns when added to the full dataset. Beside the possible biomechanical factors influencing the observed MRS trajectories, the closeness of the MRS age trajectories of <italic>Pan</italic> and modern humans, and their specificity when compared with those of <italic>Gorilla</italic> raise questions about the possible evolutionary meaning of this pattern. More specifically, it mirrors previous arguments about derived vs. shared life history traits in modern humans compared with African great apes in the literature <xref rid="pone.0107963-Zollikofer2" ref-type="bibr">[81]</xref>. Overall, our results are not consistent with a unique pattern of development and aging in modern humans. On the other hand, the possibility of parallel evolution of these traits in modern humans and <italic>Pan</italic> seems difficult to accept considering the different types of biomechanical stress (i.e. locomotory patterns) experienced by these taxa as well as the dissimilarity of other variables possibly affecting MRS development (e.g. body size, diet). Accordingly, it is plausible to postulate between modern humans and <italic>Pan</italic> a suite of synapomorphic patterns in the development of their musculoskeletal system in common with their last common ancestor.</p><p>Results on enthesopathies show: (a) an increase of EF with age only in modern humans, (b) different trajectories of OL in modern humans and African great apes, but a partial overlap between taxa of the sites showing correlation between OL and age. The lower frequencies of EF in African great apes can be compared with previous works <xref rid="pone.0107963-Jurmain2" ref-type="bibr">[53]</xref>,<xref rid="pone.0107963-Rothschild1" ref-type="bibr">[103]</xref> which showed low frequencies of degenerative joint disease in great apes. Such results, besides being possibly correlated to interspecific differences in locomotory patterns (e.g. the marked development of EF at the level of the Achilles tendon in modern humans and not in African great apes), should be considered as a function of age. The positive correlation of EF with age in modern humans and their higher frequency in elderly subjects <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>,<xref rid="pone.0107963-Milella3" ref-type="bibr">[97]</xref> indeed points to interspecific differences in maximum potential lifespan <xref rid="pone.0107963-Robson2" ref-type="bibr">[82]</xref> as the main factor underlying the observed patterns. This seems further confirmed by clinical research showing a degenerative origin of these EC <xref rid="pone.0107963-Benjamin1" ref-type="bibr">[24]</xref>,<xref rid="pone.0107963-Jiang1" ref-type="bibr">[104]</xref>,<xref rid="pone.0107963-Milz1" ref-type="bibr">[105]</xref>. Accordingly, the lack of correlation between EF and age in African great apes could be linked to a shorter lifespan accompanied by a similar (or even slower) rate of degenerative processes when compared with modern humans. Results on OL are consistent with the presence in African great apes of remodeling patterns at the level of entheses similar to what has already been observed during growth in modern humans <xref rid="pone.0107963-Mariotti2" ref-type="bibr">[18]</xref>,<xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>,<xref rid="pone.0107963-Hoyte1" ref-type="bibr">[106]</xref>–<xref rid="pone.0107963-Drfl2" ref-type="bibr">[108]</xref>. On the other hand, the increase of porosities with age in modern humans and its lack in African great apes is, as for EF, consistent with degenerative processes (see <xref rid="pone.0107963-Mariotti2" ref-type="bibr">[18]</xref>,<xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>,<xref rid="pone.0107963-Benjamin4" ref-type="bibr">[109]</xref>) specific to modern humans and mostly related to their longer lifespan.</p><p>Recent analyses of mountain gorillas <xref rid="pone.0107963-Ruff2" ref-type="bibr">[68]</xref> evidenced patterns of long bone strength consistent with ontogenetic changes in locomotory behavior. In our study, the small sample size led us to group African great apes only at the level of genus, a fact that hampers fine-grained analyses of specific ontogenetic patterns. However, the distinct age trajectories described by MRS, and the partial correlation showed between EC and long bones geometric properties <xref rid="pone.0107963-Benjamin5" ref-type="bibr">[110]</xref> make the extension of our study to the interspecific level a promising line of future research.</p></sec><sec id="s4c"><title>Sexual dimorphism</title><p>Our second hypothesis postulated higher entheseal changes in males <italic>Gorilla</italic> when compared with both modern humans and <italic>Pan</italic>. This was only partially confirmed by our results, and specifically only by sexual differences in MRS (<xref ref-type="supplementary-material" rid="pone.0107963.s013">Table S13</xref>), which evidence a scenario where <italic>Gorilla</italic> and modern humans appear markedly dimorphic when compared with <italic>Pan.</italic> Conversely, when comparing interspecific degrees of MRS sexual dimorphism, <italic>Gorilla</italic> shows a more pronounced difference between sexes. Even recognizing the possible influence on this result of the different age distribution between sexes in <italic>Gorilla</italic> (see above and <xref ref-type="table" rid="pone-0107963-t001">Table 1</xref>), our data are interestingly consistent with previous studies comparing degrees of sexual dimorphism among great apes <xref rid="pone.0107963-Wood1" ref-type="bibr">[86]</xref>. Specifically, the intermediate position of modern humans between <italic>Gorilla</italic> and <italic>Pan</italic> is consistent with the results of Lovejoy and colleagues <xref rid="pone.0107963-Lovejoy1" ref-type="bibr">[111]</xref>,<xref rid="pone.0107963-Richmond1" ref-type="bibr">[112]</xref> on articular dimensions. The higher sexual dimorphism of modern humans when compared with <italic>Pan</italic> can be traced back to a marked dimorphism in body size in the human population used in this study. EC have been indeed found to correlate with size <xref rid="pone.0107963-Weiss1" ref-type="bibr">[25]</xref>,<xref rid="pone.0107963-Weiss2" ref-type="bibr">[26]</xref>,<xref rid="pone.0107963-Niinimki3" ref-type="bibr">[113]</xref>. Note moreover that the type of sexual dimorphism observed in our dataset is consistent with previous studies on EC <xref rid="pone.0107963-Eshed1" ref-type="bibr">[6]</xref>,<xref rid="pone.0107963-Hawkey2" ref-type="bibr">[15]</xref>,<xref rid="pone.0107963-Villotte4" ref-type="bibr">[22]</xref>,<xref rid="pone.0107963-Peterson1" ref-type="bibr">[114]</xref>. As an alternative (or possible complementary) explanation, note that previous analyses on the same human dataset <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref> evidenced a marked sexual dimorphism starting from fifty years old. It is therefore possible that, as for age trajectories, the different degree of sexual dimorphism observed in modern humans and <italic>Pan</italic> is partially correlated with the longer lifespan in modern humans. An additional factor could be a marked subdivision of physical activities between the sexes in the population of Sassari. This, coupled with a longer lifespan, could have led to a higher expression of differences between sexes.</p><p>Concerning EF and OL, note that intersex comparisons in <italic>Gorilla</italic> and <italic>Pan</italic> are strongly hampered by both the small sample sizes of these taxa and the general poor expression of EF in our African great apes sample (see above). It is therefore possible that different results could be obtained by analyzing a larger dataset of both taxa. It is moreover possible that the grouping in our intersex analyses of all age classes masks patterns of sexual dimorphism more nuanced than the ones observed in MRS. Both EF and OL, when controlling for age, show complex (though not obvious) differences between sexes in modern humans <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref>,<xref rid="pone.0107963-Milella3" ref-type="bibr">[97]</xref>.</p></sec></sec><sec id="s5"><title>Conclusions</title><p>In the present study, we compared age trajectories and patterns of sexual dimorphism in entheseal robusticity and enthesopathic development between modern humans, <italic>Pan</italic> and <italic>Gorilla</italic>. Intraobserver error for the scoring of enthesopathies and MRS is overall low, pointing to a good reliability of our data.</p><p>Results revealed specific age trajectories among taxa that can be interpreted both in terms of different life history schedules and phyletic relationships. Robusticity trajectories separate <italic>Pan</italic> and modern humans from <italic>Gorilla</italic>, whereas modern humans show unique patterns of enthesopathic development which are likely the result of their extended potential lifespan. As far as sexual dimorphism is concerned, our results suggest the possible relevance of size, lifespan and physical activity in expressing differences between sexes, stressing on the other hand the need for larger samples in order to better check patterns of sexual dimorphism in enthesopathic development The present study confirmed previous hypotheses <xref rid="pone.0107963-Milella1" ref-type="bibr">[19]</xref> on the possible relevance of EC in the study of life history, pointing moreover to their usefulness in evolutionary studies.</p></sec><sec sec-type="supplementary-material" id="s6"><title>Supporting Information</title><supplementary-material content-type="local-data" id="pone.0107963.s001"><label>Table S1</label><caption><p>a) Results of the Wilcoxon test for the intraobserver error in the scoring of mean robusticty scores between two independent sessions: b) Percentage of intraobserver disagreement and Cohen’s weighted kappa for EF and OL.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s001.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s002"><label>Table S2</label><caption><p><bold>Proliferative enthesopathies (EF): summary statistics by genus and age class.</bold> SD = standard deviation.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s002.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s003"><label>Table S3</label><caption><p><bold>Osteolythic enthesopathies (OL): summary statistics by genus and age class.</bold> SD = standard deviation.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s003.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s004"><label>Table S4</label><caption><p><bold>Mean robusticity score (MRS) summary statistics by genus and age class.</bold> SD = standard deviation.</p><p>(XLSX)</p></caption><media xlink:href="pone.0107963.s004.xlsx"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s005"><label>Table S5</label><caption><p><bold><italic>Pan</italic></bold><bold>: Comparison between captive (C) and wild-shot (W) specimens in correlation between proliferative enthesopathies (EF) and age.</bold> Empty cells indicate cases for which the Spearman’s test was not possible.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s005.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s006"><label>Table S6</label><caption><p><bold><italic>Pan</italic></bold><bold>: Comparison between captive (C) and wild-shot (W) specimens in correlation between osteolithic formations (OL) and age.</bold> Empty cells indicate cases for which the Spearman’s test was not possible.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s006.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s007"><label>Table S7</label><caption><p><bold><italic>Pan</italic></bold><bold>: comparison between captive (C) and wild-shot (W) specimens in correlation between mean robusticity scores (MRS) and age.</bold></p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s007.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s008"><label>Table S8</label><caption><p><bold>Proliferative enthesopathes (EF): correlation with age.</bold> Empty cases indicate sites for which the Spearman’s test was not possible. Analyses in <italic>Gorilla</italic> and <italic>Pan</italic> include both adults and juvenile specimens. ns = not significant.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s008.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s009"><label>Table S9</label><caption><p><bold>Osteolythic enthesopathies (OL): correlation with age.</bold> Empty cases indicate sites for which the Spearman’s test was not possible. Analyses in <italic>Gorilla</italic> and <italic>Pan</italic> include both adults and juvenile specimens. ns = not significant.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s009.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s010"><label>Table S10</label><caption><p><bold>Correlation between mean robusticity scores (MRS) and age before and after the inclusion to the Gorilla and Pan samples of juvenile specimens.</bold> ns = not significant.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s010.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s011"><label>Table S11</label><caption><p><bold>Proliferative enthesopathies (EF): sex differences in the frequencies (%) of degree (2+3).</bold> F = females; M = males. Empty cases indicate sites for which the chi suqred test was not possible. ns = not significant.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s011.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s012"><label>Table S12</label><caption><p><bold>Osteolythic enthesopathies (OL): sex differences in the frequencies (%) of degrees (1) and (2+3).</bold> Empty cases indicates sites for which the chi squared test was not possible. F = females; M = males. ns = not significant.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s012.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s013"><label>Table S13</label><caption><p><bold>Sex differences (Wilcoxon test) of Mean Robusticity Scores (MRS).</bold> F = females;M = males; ns = not significant.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s013.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s014"><label>Appendix S1</label><caption><p><bold>List of human specimens used in this study.</bold> MAUB: Museum of Anthropology, University of Bologna, Italy.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s014.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s015"><label>Appendix S2</label><caption><p><bold>List of </bold><bold><italic>Gorilla</italic></bold><bold> and </bold><bold><italic>Pan</italic></bold><bold> specimens used in this study.</bold> RMCA: Royal Museum of Central Africa, Tervuren, Belgium; AIM: Anthropological Institute and Museum, University of Zurich, Switzerland.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s015.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s016"><label>Appendix S3</label><caption><p><bold>Proliferative enthesopathies (EF): complete dataset.</bold> Empty cases indicate missing data.</p><p>(XLSX)</p></caption><media xlink:href="pone.0107963.s016.xlsx"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s017"><label>Appendix S4</label><caption><p><bold>Osteolythic enthesopathies (OL): complete dataset.</bold> Empty cases indicate missing data.</p><p>(XLSX)</p></caption><media xlink:href="pone.0107963.s017.xlsx"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0107963.s018"><label>Appendix S5</label><caption><p><bold>Mean robusticity scores (MRS): complete dataset.</bold> Empty cases indicate missing data.</p><p>(XLS)</p></caption><media xlink:href="pone.0107963.s018.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material></sec></body><back><ack><p>The author wish to thank the editor and the reviewers for detailed comments, which helped improve this manuscript. Thanks to Maria Giovanna Belcastro and Valentina Mariotti for granting access to the Frassetto identified skeletal collection of Sassari at the Museum of Anthropology of the University of Bologna and for help during the data collection. Thanks to Emmanuell Gilissen and Wim Wendelen for granting access to the primatological collection of the Royal Museum of Central Africa in Tervuren. Thanks to Marcia Ponce de León for granting access to the primatological collection at the Anthropological Institute, University of Zurich. Thanks to Ann Margvelashvili for help in the scoring of dental wear. Thanks to Christoph P.E. Zollikofer, Marcia Ponce de León, John David Weissmann, Wesley Paul Petersen and Simone Callegari for useful comments on the manuscript. The present study was performed with the support of the University of Zurich.</p></ack><ref-list><title>References</title><ref id="pone.0107963-Jurmain1"><label>1</label><mixed-citation publication-type="other">Jurmain R, Villotte S (2010) Terminology. Entheses in medical literature and physical anthropology: a brief review [online]. Document published online in 4th February following the Workshop in Musculoskeletal Stress Markers (MSM): limitations and achievements in the reconstruction of past activity patterns, University of Coimbra, July 2–3, 2009. Coimbra, CIAS–Centro de Investigação em Antropologia e Saúde. Available: <ext-link ext-link-type="uri" xlink:href="http://www.uc.pt/en/cia/msm/MSM_terminology3.pdf">http://www.uc.pt/en/cia/msm/MSM_terminology3.pdf</ext-link>. Accessed 2014 Aug 27.</mixed-citation></ref><ref id="pone.0107963-Lieverse1"><label>2</label><mixed-citation publication-type="journal"><name><surname>Lieverse</surname><given-names>AR</given-names></name>, <name><surname>Bazaliiskii</surname><given-names>VI</given-names></name>, <name><surname>Goriunova</surname><given-names>OI</given-names></name>, <name><surname>Weber</surname><given-names>AW</given-names></name> (<year>2013</year>) <article-title>Lower limb activity in the Cis-Baikal: Entheseal changes among middle holocene siberian foragers</article-title>. <source>Am J Phys Anthropol</source><volume>150</volume>: <fpage>421</fpage>–<lpage>432</lpage>.<pub-id pub-id-type="pmid">23359131</pub-id></mixed-citation></ref><ref id="pone.0107963-Lieverse2"><label>3</label><mixed-citation publication-type="journal"><name><surname>Lieverse</surname><given-names>AR</given-names></name>, <name><surname>Bazaliiskii</surname><given-names>VI</given-names></name>, <name><surname>Goriunova</surname><given-names>OI</given-names></name>, <name><surname>Weber</surname><given-names>AW</given-names></name> (<year>2009</year>) <article-title>Upper limb musculoskeletal stress markers among middle Holocene foragers of Siberia’s Cis-Baikal region</article-title>. <source>Am J Phys Anthropol</source><volume>138</volume>: <fpage>458</fpage>–<lpage>472</lpage>.<pub-id pub-id-type="pmid">19085996</pub-id></mixed-citation></ref><ref id="pone.0107963-Dutour1"><label>4</label><mixed-citation publication-type="journal"><name><surname>Dutour</surname><given-names>O</given-names></name> (<year>1986</year>) <article-title>Enthesopathies (lesions of muscular insertions) as indicators of the activities of Neolithic Saharan populations</article-title>. <source>Am J Phys Anthropol</source><volume>71</volume>: <fpage>221</fpage>–<lpage>224</lpage>.<pub-id pub-id-type="pmid">3541646</pub-id></mixed-citation></ref><ref id="pone.0107963-Molnar1"><label>5</label><mixed-citation publication-type="journal"><name><surname>Molnar</surname><given-names>P</given-names></name> (<year>2005</year>) <article-title>Tracing prehistoric activities: Musculoskeletal stress marker analysis of a Stone-Age population on the island of Gotland in the Baltic Sea</article-title>. <source>Am J Phys Anthropol</source><volume>129</volume>: <fpage>12</fpage>–<lpage>23</lpage>.<pub-id pub-id-type="pmid">16161142</pub-id></mixed-citation></ref><ref id="pone.0107963-Eshed1"><label>6</label><mixed-citation publication-type="journal"><name><surname>Eshed</surname><given-names>V</given-names></name>, <name><surname>Gopher</surname><given-names>A</given-names></name>, <name><surname>Galili</surname><given-names>E</given-names></name>, <name><surname>Hershkovitz</surname><given-names>I</given-names></name> (<year>2003</year>) <article-title>Musculoskeletal stress markers in Natufian hunter-gatherers and Neolithic farmers in the Levant: The upper limb</article-title>. <source>Am J Phys Anthropol</source><volume>123</volume>: <fpage>303</fpage>–<lpage>315</lpage>.<pub-id pub-id-type="pmid">15022359</pub-id></mixed-citation></ref><ref id="pone.0107963-Molleson1"><label>7</label><mixed-citation publication-type="journal"><name><surname>Molleson</surname><given-names>T</given-names></name> (<year>1989</year>) <article-title>Seed preparation in the Mesolithic: the osteological evidence</article-title>. <source>Antiquity</source><volume>63</volume>: <fpage>356</fpage>–<lpage>362</lpage>.</mixed-citation></ref><ref id="pone.0107963-Lovell1"><label>8</label><mixed-citation publication-type="journal"><name><surname>Lovell</surname><given-names>NC</given-names></name>, <name><surname>Dublenko</surname><given-names>AA</given-names></name> (<year>1999</year>) <article-title>Further aspects of fur trade life depicted in the skeleton</article-title>. <source>Int J Osteoarch</source><volume>9</volume>: <fpage>248</fpage>–<lpage>256</lpage>.</mixed-citation></ref><ref id="pone.0107963-Henderson1"><label>9</label><mixed-citation publication-type="journal"><name><surname>Henderson</surname><given-names>C</given-names></name> (<year>2013</year>) <article-title>Subsistence strategy changes: The evidence of entheseal changes</article-title>. <source>HOMO - Journal of Comparative Human Biology</source><volume>64</volume>: <fpage>491</fpage>–<lpage>508</lpage>.<pub-id pub-id-type="pmid">24028816</pub-id></mixed-citation></ref><ref id="pone.0107963-Villotte1"><label>10</label><mixed-citation publication-type="journal"><name><surname>Villotte</surname><given-names>S</given-names></name>, <name><surname>Knusel</surname><given-names>CJ</given-names></name> (<year>2014</year>) <article-title>“I sing of arms and of a man…”: medial epicondylosis and the sexual division of labour in prehistoric Europe</article-title>. <source>J Archaeol Sci</source><volume>43</volume>: <fpage>168</fpage>–<lpage>174</lpage>.</mixed-citation></ref><ref id="pone.0107963-Hawkey1"><label>11</label><mixed-citation publication-type="journal"><name><surname>Hawkey</surname><given-names>DE</given-names></name> (<year>1998</year>) <article-title>Disability, compassion and the skeletal record: Using musculoskeletal stress markers (MSM) to construct and osteobiography from early New Mexico</article-title>. <source>Int J Osteoarch</source><volume>8</volume>: <fpage>326</fpage>–<lpage>340</lpage>.</mixed-citation></ref><ref id="pone.0107963-Havelkova1"><label>12</label><mixed-citation publication-type="journal"><name><surname>Havelkova</surname><given-names>P</given-names></name>, <name><surname>Hladik</surname><given-names>M</given-names></name>, <name><surname>Veleminsky</surname><given-names>P</given-names></name> (<year>2013</year>) <article-title>Entheseal Changes: Do They Reflect Socioeconomic Status in the Early Medieval Central European Population? (Mikulcice - Klasterisko, Great Moravian Empire, 9th–10th century)</article-title>. <source>Int J Osteoarch</source><volume>23</volume>: <fpage>237</fpage>–<lpage>251</lpage>.</mixed-citation></ref><ref id="pone.0107963-Havelkova2"><label>13</label><mixed-citation publication-type="journal"><name><surname>Havelkova</surname><given-names>P</given-names></name>, <name><surname>Villotte</surname><given-names>S</given-names></name>, <name><surname>Veleminsky</surname><given-names>P</given-names></name>, <name><surname>Polacek</surname><given-names>L</given-names></name>, <name><surname>Dobisikova</surname><given-names>M</given-names></name> (<year>2011</year>) <article-title>Enthesopathies and Activity Patterns in the Early Medieval Great Moravian Population: Evidence of Division of Labour</article-title>. <source>Int J Osteoarch</source><volume>21</volume>: <fpage>487</fpage>–<lpage>504</lpage>.</mixed-citation></ref><ref id="pone.0107963-Villotte2"><label>14</label><mixed-citation publication-type="journal"><name><surname>Villotte</surname><given-names>S</given-names></name>, <name><surname>Churchill</surname><given-names>SE</given-names></name>, <name><surname>Dutour</surname><given-names>OJ</given-names></name>, <name><surname>Henry-Gambier</surname><given-names>D</given-names></name> (<year>2010</year>) <article-title>Subsistence activities and the sexual division of labor in the European Upper Paleolithic and Mesolithic: Evidence from upper limb enthesopathies</article-title>. <source>J Hum Evol</source><volume>59</volume>: <fpage>35</fpage>–<lpage>43</lpage>.<pub-id pub-id-type="pmid">20602985</pub-id></mixed-citation></ref><ref id="pone.0107963-Hawkey2"><label>15</label><mixed-citation publication-type="journal"><name><surname>Hawkey</surname><given-names>DE</given-names></name>, <name><surname>Merbs</surname><given-names>CF</given-names></name> (<year>1995</year>) <article-title>Activity-induced musculoskeletal stress markers (MSM) and subsistence strategy changes among ancient Hudson Bay Eskimos</article-title>. <source>Int J Osteoarch</source><volume>5</volume>: <fpage>324</fpage>–<lpage>338</lpage>.</mixed-citation></ref><ref id="pone.0107963-Steen1"><label>16</label><mixed-citation publication-type="journal"><name><surname>Steen</surname><given-names>SL</given-names></name>, <name><surname>Lane</surname><given-names>RW</given-names></name> (<year>1998</year>) <article-title>Evaluation of habitual activities among two Alaskan Eskimo populations based on musculoskeletal stress markers</article-title>. <source>Int J Osteoarch</source><volume>8</volume>: <fpage>341</fpage>–<lpage>353</lpage>.</mixed-citation></ref><ref id="pone.0107963-Mariotti1"><label>17</label><mixed-citation publication-type="journal"><name><surname>Mariotti</surname><given-names>V</given-names></name>, <name><surname>Facchini</surname><given-names>F</given-names></name>, <name><surname>Belcastro</surname><given-names>MG</given-names></name> (<year>2007</year>) <article-title>The study of entheses: proposal of a standardised scoring method for twenty-three entheses of the postcranial skeleton</article-title>. <source>Coll Antropol</source><volume>31</volume>: <fpage>291</fpage>–<lpage>313</lpage>.<pub-id pub-id-type="pmid">17598416</pub-id></mixed-citation></ref><ref id="pone.0107963-Mariotti2"><label>18</label><mixed-citation publication-type="journal"><name><surname>Mariotti</surname><given-names>V</given-names></name>, <name><surname>Facchini</surname><given-names>F</given-names></name>, <name><surname>Belcastro</surname><given-names>MG</given-names></name> (<year>2004</year>) <article-title>Enthesopathies–proposal of a standardized scoring method and applications</article-title>. <source>Coll Antropol</source><volume>28</volume>: <fpage>145</fpage>–<lpage>159</lpage>.<pub-id pub-id-type="pmid">15636072</pub-id></mixed-citation></ref><ref id="pone.0107963-Milella1"><label>19</label><mixed-citation publication-type="journal"><name><surname>Milella</surname><given-names>M</given-names></name>, <name><surname>Belcastro</surname><given-names>MG</given-names></name>, <name><surname>Zollikofer</surname><given-names>CP</given-names></name>, <name><surname>Mariotti</surname><given-names>V</given-names></name> (<year>2012</year>) <article-title>The effect of age, sex, and physical activity on entheseal morphology in a contemporary Italian skeletal collection</article-title>. <source>Am J Phys Anthropol</source><volume>148</volume>: <fpage>379</fpage>–<lpage>388</lpage>.<pub-id pub-id-type="pmid">22460619</pub-id></mixed-citation></ref><ref id="pone.0107963-AlvesCardoso1"><label>20</label><mixed-citation publication-type="journal"><name><surname>Alves Cardoso</surname><given-names>F</given-names></name>, <name><surname>Henderson</surname><given-names>CY</given-names></name> (<year>2010</year>) <article-title>Enthesopathy formation in the humerus: Data from known age at death and known occupation skeletal collections</article-title>. <source>Am J Phys Anthropol</source><volume>141</volume>: <fpage>550</fpage>–<lpage>560</lpage>.<pub-id pub-id-type="pmid">19927279</pub-id></mixed-citation></ref><ref id="pone.0107963-Villotte3"><label>21</label><mixed-citation publication-type="journal"><name><surname>Villotte</surname><given-names>S</given-names></name>, <name><surname>Castex</surname><given-names>D</given-names></name>, <name><surname>Couallier</surname><given-names>V</given-names></name>, <name><surname>Dutour</surname><given-names>O</given-names></name>, <name><surname>Knüsel</surname><given-names>CJ</given-names></name>, <etal>et al</etal> (<year>2010</year>) <article-title>Enthesopathies as occupational stress markers: evidence from the upper limb</article-title>. <source>Am J Phys Anthropol</source><volume>142</volume>: <fpage>224</fpage>–<lpage>234</lpage>.<pub-id pub-id-type="pmid">20034011</pub-id></mixed-citation></ref><ref id="pone.0107963-Villotte4"><label>22</label><mixed-citation publication-type="other">Villotte S (2009) Enthésopathies et activités des hommes préhistoriques: recherche méthodologique et application aux fossiles européens du Paléolithique supérieur et du Mésolithique: Archaeopress.</mixed-citation></ref><ref id="pone.0107963-Niinimki1"><label>23</label><mixed-citation publication-type="journal"><name><surname>Niinimäki</surname><given-names>S</given-names></name> (<year>2009</year>) <article-title>What do muscle marker ruggedness scores actually tell us?</article-title><source>Int J Osteoarch</source><volume>21</volume>: <fpage>292</fpage>–<lpage>299</lpage>.</mixed-citation></ref><ref id="pone.0107963-Benjamin1"><label>24</label><mixed-citation publication-type="journal"><name><surname>Benjamin</surname><given-names>M</given-names></name>, <name><surname>Toumi</surname><given-names>H</given-names></name>, <name><surname>Suzuki</surname><given-names>D</given-names></name>, <name><surname>Hayashi</surname><given-names>K</given-names></name>, <name><surname>McGonagle</surname><given-names>D</given-names></name> (<year>2009</year>) <article-title>Evidence for a distinctive pattern of bone formation in enthesophytes</article-title>. <source>Ann Rheum Dis</source><volume>68</volume>: <fpage>1003</fpage>–<lpage>1010</lpage>.<pub-id pub-id-type="pmid">18625624</pub-id></mixed-citation></ref><ref id="pone.0107963-Weiss1"><label>25</label><mixed-citation publication-type="journal"><name><surname>Weiss</surname><given-names>E</given-names></name> (<year>2003</year>) <article-title>Understanding muscle markers: aggregation and construct validity</article-title>. <source>Am J Phys Anthropol</source><volume>121</volume>: <fpage>230</fpage>–<lpage>240</lpage>.<pub-id pub-id-type="pmid">12772211</pub-id></mixed-citation></ref><ref id="pone.0107963-Weiss2"><label>26</label><mixed-citation publication-type="journal"><name><surname>Weiss</surname><given-names>E</given-names></name> (<year>2004</year>) <article-title>Understanding muscle markers: lower limbs</article-title>. <source>Am J Phys Anthropol</source><volume>125</volume>: <fpage>232</fpage>–<lpage>238</lpage>.<pub-id pub-id-type="pmid">15386252</pub-id></mixed-citation></ref><ref id="pone.0107963-Villotte5"><label>27</label><mixed-citation publication-type="journal"><name><surname>Villotte</surname><given-names>S</given-names></name>, <name><surname>Knüsel</surname><given-names>CJ</given-names></name> (<year>2014</year>) <article-title>“I sing of arms and of a man…”: medial epicondylosis and the sexual division of labour in prehistoric Europe</article-title>. <source>J Archaeol Sci</source><volume>43</volume>: <fpage>168</fpage>–<lpage>174</lpage>.</mixed-citation></ref><ref id="pone.0107963-alOumaoui1"><label>28</label><mixed-citation publication-type="journal"><name><surname>al-Oumaoui</surname><given-names>I</given-names></name>, <name><surname>Jimenez-Brobeil</surname><given-names>S</given-names></name>, <name><surname>du Souich</surname><given-names>P</given-names></name> (<year>2004</year>) <article-title>Markers of activity patterns in some populations of the Iberian Peninsula</article-title>. <source>Int J Osteoarch</source><volume>14</volume>: <fpage>343</fpage>–<lpage>359</lpage>.</mixed-citation></ref><ref id="pone.0107963-Wilczak1"><label>29</label><mixed-citation publication-type="journal"><name><surname>Wilczak</surname><given-names>CA</given-names></name> (<year>1998</year>) <article-title>Consideration of sexual dimorphism, age, and asymmetry in quantitative measurements of muscle insertion sites</article-title>. <source>Int J Osteoarch</source><volume>8</volume>: <fpage>311</fpage>–<lpage>325</lpage>.</mixed-citation></ref><ref id="pone.0107963-Benjamin2"><label>30</label><mixed-citation publication-type="journal"><name><surname>Benjamin</surname><given-names>M</given-names></name>, <name><surname>Kumai</surname><given-names>T</given-names></name>, <name><surname>Milz</surname><given-names>S</given-names></name>, <name><surname>Boszczyk</surname><given-names>BM</given-names></name>, <name><surname>Boszczyk</surname><given-names>AA</given-names></name>, <etal>et al</etal> (<year>2002</year>) <article-title>The skeletal attachment of tendons–tendon “entheses”</article-title>. <source>Comp Biochem Physiol A Mol Integr Physiol</source><volume>133</volume>: <fpage>931</fpage>–<lpage>945</lpage>.<pub-id pub-id-type="pmid">12485684</pub-id></mixed-citation></ref><ref id="pone.0107963-Benjamin3"><label>31</label><mixed-citation publication-type="journal"><name><surname>Benjamin</surname><given-names>M</given-names></name>, <name><surname>Ralphs</surname><given-names>JR</given-names></name> (<year>1995</year>) <article-title>Functional and developmental anatomy of tendons and ligaments</article-title>. <source>Repetitive motion disorders of the upper extremity Rosemont, IL: American Academy of Orthopaedic Surgeons</source><volume>1995</volume>: <fpage>185</fpage>–<lpage>203</lpage>.</mixed-citation></ref><ref id="pone.0107963-Villotte6"><label>32</label><mixed-citation publication-type="other">Villotte S (2006) Connaissances médicales actuelles, cotation des enthésopathies: nouvelle méthode. Bulletins et Mémoires de la Société d’Anthropologie de Paris: 65–85.</mixed-citation></ref><ref id="pone.0107963-Henderson2"><label>33</label><mixed-citation publication-type="journal"><name><surname>Henderson</surname><given-names>CY</given-names></name>, <name><surname>Mariotti</surname><given-names>V</given-names></name>, <name><surname>Pany-Kucera</surname><given-names>D</given-names></name>, <name><surname>Villotte</surname><given-names>S</given-names></name>, <name><surname>Wilczak</surname><given-names>C</given-names></name> (<year>2013</year>) <article-title>Recording Specific Entheseal Changes of Fibrocartilaginous Entheses: Initial Tests Using the Coimbra Method</article-title>. <source>Int J Osteoarch</source><volume>23</volume>: <fpage>152</fpage>–<lpage>162</lpage>.</mixed-citation></ref><ref id="pone.0107963-Durigon1"><label>34</label><mixed-citation publication-type="book">Durigon M, Paolaggi J (1991) Enthèse au cours de la vie. Pathologie des insertions et enthésopathies Paris: Masson p: 12–17.</mixed-citation></ref><ref id="pone.0107963-Robb1"><label>35</label><mixed-citation publication-type="journal"><name><surname>Robb</surname><given-names>JE</given-names></name> (<year>1998</year>) <article-title>The interpretation of skeletal muscle sites: A statistical approach</article-title>. <source>Int J Osteoarch</source><volume>8</volume>: <fpage>363</fpage>–<lpage>377</lpage>.</mixed-citation></ref><ref id="pone.0107963-Milella2"><label>36</label><mixed-citation publication-type="journal"><name><surname>Milella</surname><given-names>M</given-names></name>, <name><surname>Zollikofer</surname><given-names>CP</given-names></name>, <name><surname>De León</surname><given-names>MP</given-names></name> (<year>2013</year>) <article-title>Patterns of entheseal changes in modern humans and African great apes</article-title>. <source>Am J Phys Anthropol</source><volume>150</volume>: <fpage>198</fpage>.</mixed-citation></ref><ref id="pone.0107963-Morimoto1"><label>37</label><mixed-citation publication-type="journal"><name><surname>Morimoto</surname><given-names>N</given-names></name>, <name><surname>Zollikofer</surname><given-names>CP</given-names></name>, <name><surname>Ponce de Leon</surname><given-names>MS</given-names></name> (<year>2012</year>) <article-title>Shared human-chimpanzee pattern of perinatal femoral shaft morphology and its implications for the evolution of hominin locomotor adaptations</article-title>. <source>PLOS One</source><volume>7</volume>: <fpage>e41980</fpage>.<pub-id pub-id-type="pmid">22848680</pub-id></mixed-citation></ref><ref id="pone.0107963-Morimoto2"><label>38</label><mixed-citation publication-type="journal"><name><surname>Morimoto</surname><given-names>N</given-names></name>, <name><surname>Ponce de Leon</surname><given-names>MS</given-names></name>, <name><surname>Zollikofer</surname><given-names>CP</given-names></name> (<year>2011</year>) <article-title>Exploring femoral diaphyseal shape variation in wild and captive chimpanzees by means of morphometric mapping: a test of Wolff’s law</article-title>. <source>Anat Rec (Hoboken)</source><volume>294</volume>: <fpage>589</fpage>–<lpage>609</lpage>.<pub-id pub-id-type="pmid">21328564</pub-id></mixed-citation></ref><ref id="pone.0107963-Maggiano1"><label>39</label><mixed-citation publication-type="journal"><name><surname>Maggiano</surname><given-names>IS</given-names></name>, <name><surname>Schultz</surname><given-names>M</given-names></name>, <name><surname>Kierdorf</surname><given-names>H</given-names></name>, <name><surname>Sosa</surname><given-names>TS</given-names></name>, <name><surname>Maggiano</surname><given-names>CM</given-names></name>, <etal>et al</etal> (<year>2008</year>) <article-title>Cross-sectional analysis of long bones, occupational activities and long-distance trade of the Classic Maya from Xcambo–archaeological and osteological evidence</article-title>. <source>Am J Phys Anthropol</source><volume>136</volume>: <fpage>470</fpage>–<lpage>477</lpage>.<pub-id pub-id-type="pmid">18383157</pub-id></mixed-citation></ref><ref id="pone.0107963-Marchi1"><label>40</label><mixed-citation publication-type="journal"><name><surname>Marchi</surname><given-names>D</given-names></name> (<year>2008</year>) <article-title>Relationships between lower limb cross-sectional geometry and mobility: The case of a Neolithic sample from Italy</article-title>. <source>Am J Phys Anthropol</source><volume>137</volume>: <fpage>188</fpage>–<lpage>200</lpage>.<pub-id pub-id-type="pmid">18470890</pub-id></mixed-citation></ref><ref id="pone.0107963-Marchi2"><label>41</label><mixed-citation publication-type="journal"><name><surname>Marchi</surname><given-names>D</given-names></name> (<year>2010</year>) <article-title>Articular to diaphyseal proportions of human and great ape metatarsals</article-title>. <source>Am J Phys Anthropol</source><volume>143</volume>: <fpage>198</fpage>–<lpage>207</lpage>.<pub-id pub-id-type="pmid">20853475</pub-id></mixed-citation></ref><ref id="pone.0107963-Patel1"><label>42</label><mixed-citation publication-type="journal"><name><surname>Patel</surname><given-names>BA</given-names></name>, <name><surname>Ruff</surname><given-names>CB</given-names></name>, <name><surname>Simons</surname><given-names>EL</given-names></name>, <name><surname>Organ</surname><given-names>JM</given-names></name> (<year>2013</year>) <article-title>Humeral cross-sectional shape in suspensory primates and sloths</article-title>. <source>Anat Rec (Hoboken)</source><volume>296</volume>: <fpage>545</fpage>–<lpage>556</lpage>.<pub-id pub-id-type="pmid">23408647</pub-id></mixed-citation></ref><ref id="pone.0107963-Ruff1"><label>43</label><mixed-citation publication-type="journal"><name><surname>Ruff</surname><given-names>CB</given-names></name>, <name><surname>Runestad</surname><given-names>JA</given-names></name> (<year>1992</year>) <article-title>Primate Limb Bone Structural Adaptations</article-title>. <source>Annual Review of Anthropology</source><volume>21</volume>: <fpage>407</fpage>–<lpage>433</lpage>.</mixed-citation></ref><ref id="pone.0107963-Shaw1"><label>44</label><mixed-citation publication-type="journal"><name><surname>Shaw</surname><given-names>CN</given-names></name>, <name><surname>Stock</surname><given-names>JT</given-names></name> (<year>2011</year>) <article-title>The influence of body proportions on femoral and tibial midshaft shape in hunter-gatherers</article-title>. <source>Am J Phys Anthropol</source><volume>144</volume>: <fpage>22</fpage>–<lpage>29</lpage>.<pub-id pub-id-type="pmid">20623683</pub-id></mixed-citation></ref><ref id="pone.0107963-Barak1"><label>45</label><mixed-citation publication-type="journal"><name><surname>Barak</surname><given-names>MM</given-names></name>, <name><surname>Lieberman</surname><given-names>DE</given-names></name>, <name><surname>Raichlen</surname><given-names>D</given-names></name>, <name><surname>Pontzer</surname><given-names>H</given-names></name>, <name><surname>Warrener</surname><given-names>AG</given-names></name>, <etal>et al</etal> (<year>2013</year>) <article-title>Trabecular Evidence for a Human-Like Gait in Australopithecus africanus</article-title>. <source>PLOS One</source><volume>8</volume>: <fpage>e77687</fpage>.<pub-id pub-id-type="pmid">24223719</pub-id></mixed-citation></ref><ref id="pone.0107963-Ryan1"><label>46</label><mixed-citation publication-type="journal"><name><surname>Ryan</surname><given-names>TM</given-names></name>, <name><surname>Shaw</surname><given-names>CN</given-names></name> (<year>2012</year>) <article-title>Unique suites of trabecular bone features characterize locomotor behavior in human and non-human anthropoid primates</article-title>. <source>PLOS One</source><volume>7</volume>: <fpage>e41037</fpage>.<pub-id pub-id-type="pmid">22815902</pub-id></mixed-citation></ref><ref id="pone.0107963-Shaw2"><label>47</label><mixed-citation publication-type="journal"><name><surname>Shaw</surname><given-names>CN</given-names></name>, <name><surname>Ryan</surname><given-names>TM</given-names></name> (<year>2012</year>) <article-title>Does skeletal anatomy reflect adaptation to locomotor patterns? Cortical and trabecular architecture in human and nonhuman anthropoids</article-title>. <source>American journal of physical anthropology</source><volume>147</volume>: <fpage>187</fpage>–<lpage>200</lpage>.<pub-id pub-id-type="pmid">22120605</pub-id></mixed-citation></ref><ref id="pone.0107963-Tsegai1"><label>48</label><mixed-citation publication-type="journal"><name><surname>Tsegai</surname><given-names>ZJ</given-names></name>, <name><surname>Kivell</surname><given-names>TL</given-names></name>, <name><surname>Gross</surname><given-names>T</given-names></name>, <name><surname>Nguyen</surname><given-names>NH</given-names></name>, <name><surname>Pahr</surname><given-names>DH</given-names></name>, <etal>et al</etal> (<year>2013</year>) <article-title>Trabecular bone structure correlates with hand posture and use in hominoids</article-title>. <source>PLOS One</source><volume>8</volume>: <fpage>e78781</fpage>.<pub-id pub-id-type="pmid">24244359</pub-id></mixed-citation></ref><ref id="pone.0107963-DeSilva1"><label>49</label><mixed-citation publication-type="journal"><name><surname>DeSilva</surname><given-names>JM</given-names></name>, <name><surname>Devlin</surname><given-names>MJ</given-names></name> (<year>2012</year>) <article-title>A comparative study of the trabecular bony architecture of the talus in humans, non-human primates, and Australopithecus</article-title>. <source>J Hum Evol</source><volume>63</volume>: <fpage>536</fpage>–<lpage>551</lpage>.<pub-id pub-id-type="pmid">22840715</pub-id></mixed-citation></ref><ref id="pone.0107963-Morimoto3"><label>50</label><mixed-citation publication-type="journal"><name><surname>Morimoto</surname><given-names>N</given-names></name>, <name><surname>De Leon</surname><given-names>MS</given-names></name>, <name><surname>Nishimura</surname><given-names>T</given-names></name>, <name><surname>Zollikofer</surname><given-names>CP</given-names></name> (<year>2011</year>) <article-title>Femoral Morphology and Femoropelvic Musculoskeletal Anatomy of Humans and Great Apes: A Comparative Virtopsy Study</article-title>. <source>Anat Rec (Hoboken)</source><volume>294</volume>: <fpage>1433</fpage>–<lpage>1445</lpage>.<pub-id pub-id-type="pmid">21809454</pub-id></mixed-citation></ref><ref id="pone.0107963-Kikuchi1"><label>51</label><mixed-citation publication-type="journal"><name><surname>Kikuchi</surname><given-names>Y</given-names></name>, <name><surname>Takemoto</surname><given-names>H</given-names></name>, <name><surname>Kuraoka</surname><given-names>A</given-names></name> (<year>2012</year>) <article-title>Relationship between humeral geometry and shoulder muscle power among suspensory, knuckle-walking, and digitigrade/palmigrade quadrupedal primates</article-title>. <source>J Anat</source><volume>220</volume>: <fpage>29</fpage>–<lpage>41</lpage>.<pub-id pub-id-type="pmid">22050714</pub-id></mixed-citation></ref><ref id="pone.0107963-Myatt1"><label>52</label><mixed-citation publication-type="journal"><name><surname>Myatt</surname><given-names>JP</given-names></name>, <name><surname>Crompton</surname><given-names>RH</given-names></name>, <name><surname>Payne-Davis</surname><given-names>RC</given-names></name>, <name><surname>Vereecke</surname><given-names>EE</given-names></name>, <name><surname>Isler</surname><given-names>K</given-names></name>, <etal>et al</etal> (<year>2012</year>) <article-title>Functional adaptations in the forelimb muscles of non-human great apes</article-title>. <source>J Anat</source><volume>220</volume>: <fpage>13</fpage>–<lpage>28</lpage>.<pub-id pub-id-type="pmid">22034995</pub-id></mixed-citation></ref><ref id="pone.0107963-Jurmain2"><label>53</label><mixed-citation publication-type="journal"><name><surname>Jurmain</surname><given-names>R</given-names></name> (<year>2000</year>) <article-title>Degenerative joint disease in African great apes: an evolutionary perspective</article-title>. <source>J Hum Evol</source><volume>39</volume>: <fpage>185</fpage>–<lpage>203</lpage>.<pub-id pub-id-type="pmid">10968928</pub-id></mixed-citation></ref><ref id="pone.0107963-Jurmain3"><label>54</label><mixed-citation publication-type="journal"><name><surname>Jurmain</surname><given-names>R</given-names></name> (<year>1989</year>) <article-title>Trauma, degenerative disease, and other pathologies among the Gombe chimpanzees</article-title>. <source>Am J Phys Anthropol</source><volume>80</volume>: <fpage>229</fpage>–<lpage>237</lpage>.<pub-id pub-id-type="pmid">2801915</pub-id></mixed-citation></ref><ref id="pone.0107963-Schultz1"><label>55</label><mixed-citation publication-type="journal"><name><surname>Schultz</surname><given-names>AH</given-names></name> (<year>1939</year>) <article-title>Notes on Diseases and Healed Fractures of Wild Apes and their bearing on the antiquity of pathological conditions in man</article-title>. <source>Bull Hist Med</source><volume>7</volume>: <fpage>581</fpage>–<lpage>582</lpage>.</mixed-citation></ref><ref id="pone.0107963-Schultz2"><label>56</label><mixed-citation publication-type="journal"><name><surname>Schultz</surname><given-names>AH</given-names></name> (<year>1956</year>) <article-title>The occurrence and frequency of pathological and teratological conditions and of twinning among non-human primates</article-title>. <source>Primatologia</source><volume>1</volume>: <fpage>965</fpage>–<lpage>1014</lpage>.</mixed-citation></ref><ref id="pone.0107963-Drapeau1"><label>57</label><mixed-citation publication-type="journal"><name><surname>Drapeau</surname><given-names>MS</given-names></name> (<year>2008</year>) <article-title>Enthesis bilateral asymmetry in humans and African apes</article-title>. <source>Homo: internationale Zeitschrift fur die vergleichende Forschung am Menschen</source><volume>59</volume>: <fpage>93</fpage>–<lpage>109</lpage>.<pub-id pub-id-type="pmid">18396287</pub-id></mixed-citation></ref><ref id="pone.0107963-Kelley1"><label>58</label><mixed-citation publication-type="journal"><name><surname>Kelley</surname><given-names>J</given-names></name>, <name><surname>Schwartz</surname><given-names>GT</given-names></name> (<year>2010</year>) <article-title>Dental development and life history in living African and Asian apes</article-title>. <source>Proc Natl Acad Sci USA</source><volume>107</volume>: <fpage>1035</fpage>–<lpage>1040</lpage>.<pub-id pub-id-type="pmid">20080537</pub-id></mixed-citation></ref><ref id="pone.0107963-Dean1"><label>59</label><mixed-citation publication-type="journal"><name><surname>Dean</surname><given-names>MC</given-names></name> (<year>2010</year>) <article-title>Retrieving chronological age from dental remains of early fossil hominins to reconstruct human growth in the past</article-title>. <source>Philos Trans R Soc Lond B Biol Sci</source><volume>365</volume>: <fpage>3397</fpage>–<lpage>3410</lpage>.<pub-id pub-id-type="pmid">20855313</pub-id></mixed-citation></ref><ref id="pone.0107963-Schwartz1"><label>60</label><mixed-citation publication-type="journal"><name><surname>Schwartz</surname><given-names>GT</given-names></name> (<year>2012</year>) <article-title>Growth, Development, and Life History throughout the Evolution of Homo</article-title>. <source>Curr Anthropol</source><volume>53</volume>: <fpage>S395</fpage>–<lpage>S408</lpage>.</mixed-citation></ref><ref id="pone.0107963-Zollikofer1"><label>61</label><mixed-citation publication-type="journal"><name><surname>Zollikofer</surname><given-names>CP</given-names></name> (<year>2012</year>) <article-title>Evolution of hominin cranial ontogeny</article-title>. <source>Prog Brain Res</source><volume>195</volume>: <fpage>273</fpage>.<pub-id pub-id-type="pmid">22230632</pub-id></mixed-citation></ref><ref id="pone.0107963-Lieberman1"><label>62</label><mixed-citation publication-type="journal"><name><surname>Lieberman</surname><given-names>DE</given-names></name>, <name><surname>Carlo</surname><given-names>J</given-names></name>, <name><surname>Ponce de León</surname><given-names>M</given-names></name>, <name><surname>Zollikofer</surname><given-names>CPE</given-names></name> (<year>2007</year>) <article-title>A geometric morphometric analysis of heterochrony in the cranium of chimpanzees and bonobos</article-title>. <source>J Hum Evol</source><volume>52</volume>: <fpage>647</fpage>–<lpage>662</lpage>.<pub-id pub-id-type="pmid">17298840</pub-id></mixed-citation></ref><ref id="pone.0107963-Morimoto4"><label>63</label><mixed-citation publication-type="journal"><name><surname>Morimoto</surname><given-names>N</given-names></name>, <name><surname>Ogihara</surname><given-names>N</given-names></name>, <name><surname>Katayama</surname><given-names>K</given-names></name>, <name><surname>Shiota</surname><given-names>K</given-names></name> (<year>2008</year>) <article-title>Three-dimensional ontogenetic shape changes in the human cranium during the fetal period</article-title>. <source>J Anat</source><volume>212</volume>: <fpage>627</fpage>–<lpage>635</lpage>.<pub-id pub-id-type="pmid">18430090</pub-id></mixed-citation></ref><ref id="pone.0107963-Bolter1"><label>64</label><mixed-citation publication-type="journal"><name><surname>Bolter</surname><given-names>DR</given-names></name>, <name><surname>Zihlman</surname><given-names>AL</given-names></name> (<year>2012</year>) <article-title>Skeletal development in Pan paniscus with comparisons to Pan troglodytes</article-title>. <source>Am J Phys Anthropol</source><volume>147</volume>: <fpage>629</fpage>–<lpage>636</lpage>.<pub-id pub-id-type="pmid">22331605</pub-id></mixed-citation></ref><ref id="pone.0107963-Watts1"><label>65</label><mixed-citation publication-type="journal"><name><surname>Watts</surname><given-names>E</given-names></name> (<year>1993</year>) <article-title>Epiphyseal union in captive chimpanzees</article-title>. <source>Am J Phys Anthropol</source><volume>16</volume>: <fpage>206</fpage>.</mixed-citation></ref><ref id="pone.0107963-Schultz3"><label>66</label><mixed-citation publication-type="journal"><name><surname>Schultz</surname><given-names>AH</given-names></name> (<year>1940</year>) <article-title>Growth and development of the chimpanzee</article-title>. <source>Contrib Embryol</source><volume>28</volume>: <fpage>1</fpage>–<lpage>63</lpage>.</mixed-citation></ref><ref id="pone.0107963-Zihlman1"><label>67</label><mixed-citation publication-type="journal"><name><surname>Zihlman</surname><given-names>AL</given-names></name>, <name><surname>Bolter</surname><given-names>DR</given-names></name>, <name><surname>Boesch</surname><given-names>C</given-names></name> (<year>2007</year>) <article-title>Skeletal and dental growth and development in chimpanzees of the Taï National Park, Côte D’Ivoire</article-title>. <source>J Zool</source><volume>273</volume>: <fpage>63</fpage>–<lpage>73</lpage>.</mixed-citation></ref><ref id="pone.0107963-Ruff2"><label>68</label><mixed-citation publication-type="journal"><name><surname>Ruff</surname><given-names>CB</given-names></name>, <name><surname>Burgess</surname><given-names>ML</given-names></name>, <name><surname>Bromage</surname><given-names>TG</given-names></name>, <name><surname>Mudakikwa</surname><given-names>A</given-names></name>, <name><surname>McFarlin</surname><given-names>SC</given-names></name> (<year>2013</year>) <article-title>Ontogenetic changes in limb bone structural proportions in mountain gorillas (Gorilla beringei beringei)</article-title>. <source>J Hum Evol</source><volume>65</volume>: <fpage>693</fpage>–<lpage>703</lpage>.<pub-id pub-id-type="pmid">24129040</pub-id></mixed-citation></ref><ref id="pone.0107963-DeRousseau1"><label>69</label><mixed-citation publication-type="journal"><name><surname>DeRousseau</surname><given-names>CJ</given-names></name> (<year>1985</year>) <article-title>Aging in the musculoskeletal system of rhesus monkeys: II. Degenerative joint disease</article-title>. <source>Am J Phys Anthropol</source><volume>67</volume>: <fpage>177</fpage>–<lpage>184</lpage>.<pub-id pub-id-type="pmid">4061576</pub-id></mixed-citation></ref><ref id="pone.0107963-DeRousseau2"><label>70</label><mixed-citation publication-type="journal"><name><surname>DeRousseau</surname><given-names>CJ</given-names></name> (<year>2005</year>) <article-title>Aging in the musculoskeletal system of rhesus monkeys: III. Bone loss</article-title>. <source>Am J Phys Anthropol</source><volume>68</volume>: <fpage>157</fpage>–<lpage>167</lpage>.<pub-id pub-id-type="pmid">4061606</pub-id></mixed-citation></ref><ref id="pone.0107963-DeRousseau3"><label>71</label><mixed-citation publication-type="journal"><name><surname>DeRousseau</surname><given-names>CJ</given-names></name>, <name><surname>Rawlins</surname><given-names>RG</given-names></name>, <name><surname>Denlinger</surname><given-names>JL</given-names></name> (<year>2005</year>) <article-title>Aging in the musculoskeletal system of rhesus monkeys: I. Passive joint excursion</article-title>. <source>Am J Phys Anthropol</source><volume>61</volume>: <fpage>483</fpage>–<lpage>494</lpage>.<pub-id pub-id-type="pmid">6624892</pub-id></mixed-citation></ref><ref id="pone.0107963-Pope1"><label>72</label><mixed-citation publication-type="journal"><name><surname>Pope</surname><given-names>NS</given-names></name>, <name><surname>Gould</surname><given-names>KG</given-names></name>, <name><surname>Anderson</surname><given-names>DC</given-names></name>, <name><surname>Mann</surname><given-names>DR</given-names></name> (<year>1989</year>) <article-title>Effects of age and sex on bone density in the rhesus monkey</article-title>. <source>Bone</source><volume>10</volume>: <fpage>109</fpage>–<lpage>112</lpage>.<pub-id pub-id-type="pmid">2765308</pub-id></mixed-citation></ref><ref id="pone.0107963-Sumner1"><label>73</label><mixed-citation publication-type="journal"><name><surname>Sumner</surname><given-names>DR</given-names></name>, <name><surname>Morbeck</surname><given-names>ME</given-names></name>, <name><surname>Lobick</surname><given-names>JJ</given-names></name> (<year>1989</year>) <article-title>Apparent age-related bone loss among adult female Gombe chimpanzees</article-title>. <source>Am J Phys Anthropol</source><volume>79</volume>: <fpage>225</fpage>–<lpage>234</lpage>.<pub-id pub-id-type="pmid">2742005</pub-id></mixed-citation></ref><ref id="pone.0107963-Morbeck1"><label>74</label><mixed-citation publication-type="journal"><name><surname>Morbeck</surname><given-names>ME</given-names></name>, <name><surname>Galloway</surname><given-names>A</given-names></name>, <name><surname>Sumner</surname><given-names>DR</given-names></name> (<year>2002</year>) <article-title>Getting old at Gombe: skeletal aging in wild-ranging chimpanzees</article-title>. <source>Interdisciplinary topics in gerontology</source><volume>31</volume>: <fpage>48</fpage>–<lpage>62</lpage>.</mixed-citation></ref><ref id="pone.0107963-Schultz4"><label>75</label><mixed-citation publication-type="journal"><name><surname>Schultz</surname><given-names>AH</given-names></name> (<year>1944</year>) <article-title>Age changes and variability in gibbons. A Morphological study on a population sample of a man-like ape</article-title>. <source>Am J Phys Anthropol</source><volume>2</volume>: <fpage>1</fpage>–<lpage>129</lpage>.</mixed-citation></ref><ref id="pone.0107963-vanSchaik1"><label>76</label><mixed-citation publication-type="book">van Schaik CP, Barrickman N, Bastian ML, Krakauer EB, van Noordwijk MA (2006) Primate life histories and the role of brains. In Hawkes K, Paine RR (eds) The Evolution of Human Life History. School of American Research Press, Santa Fe, NM: 127–154.</mixed-citation></ref><ref id="pone.0107963-Robson1"><label>77</label><mixed-citation publication-type="journal"><name><surname>Robson</surname><given-names>SL</given-names></name>, <name><surname>Wood</surname><given-names>B</given-names></name> (<year>2008</year>) <article-title>Hominin life history: reconstruction and evolution</article-title>. <source>J Anat</source><volume>212</volume>: <fpage>394</fpage>–<lpage>425</lpage>.<pub-id pub-id-type="pmid">18380863</pub-id></mixed-citation></ref><ref id="pone.0107963-Karasik1"><label>78</label><mixed-citation publication-type="journal"><name><surname>Karasik</surname><given-names>D</given-names></name>, <name><surname>Kiel</surname><given-names>DP</given-names></name> (<year>2010</year>) <article-title>Evidence for pleiotropic factors in genetics of the musculoskeletal system</article-title>. <source>Bone</source><volume>46</volume>: <fpage>1226</fpage>–<lpage>1237</lpage>.<pub-id pub-id-type="pmid">20149904</pub-id></mixed-citation></ref><ref id="pone.0107963-Bogin1"><label>79</label><mixed-citation publication-type="journal"><name><surname>Bogin</surname><given-names>BA</given-names></name>, <name><surname>Smith</surname><given-names>BH</given-names></name> (<year>1996</year>) <article-title>Evolution of the human life cycle</article-title>. <source>Am J Hum Biol</source><volume>8</volume>: <fpage>703</fpage>–<lpage>716</lpage>.</mixed-citation></ref><ref id="pone.0107963-BlurtonJones1"><label>80</label><mixed-citation publication-type="other">Blurton Jones N (2006) Contemporary hunter-gatherers and human life history evolution. In Hawkes K, Paine RR (eds) The evolution of human life history: 231–266.</mixed-citation></ref><ref id="pone.0107963-Zollikofer2"><label>81</label><mixed-citation publication-type="journal"><name><surname>Zollikofer</surname><given-names>CPE</given-names></name>, <name><surname>Ponce de León</surname><given-names>MS</given-names></name> (<year>2010</year>) <article-title>The evolution of hominin ontogenies</article-title>. <source>Semin Cell Dev Biol</source><volume>21</volume>: <fpage>441</fpage>–<lpage>452</lpage>.<pub-id pub-id-type="pmid">19900572</pub-id></mixed-citation></ref><ref id="pone.0107963-Robson2"><label>82</label><mixed-citation publication-type="book">Robson SL, Van Schaik CP, Hawkes K (2006) The derived features of human life history. In Hawkes K, Paine RR (eds) The evolution of human life history. School of American Research Press, Santa Fe, NM: 17–44.</mixed-citation></ref><ref id="pone.0107963-Kaplan1"><label>83</label><mixed-citation publication-type="journal"><name><surname>Kaplan</surname><given-names>H</given-names></name>, <name><surname>Hill</surname><given-names>K</given-names></name>, <name><surname>Lancaster</surname><given-names>J</given-names></name>, <name><surname>Hurtado</surname><given-names>AM</given-names></name> (<year>2000</year>) <article-title>A theory of human life history evolution: diet, intelligence, and longevity</article-title>. <source>Evol Anthropol</source><volume>9</volume>: <fpage>156</fpage>–<lpage>185</lpage>.</mixed-citation></ref><ref id="pone.0107963-Leigh1"><label>84</label><mixed-citation publication-type="journal"><name><surname>Leigh</surname><given-names>SR</given-names></name> (<year>1996</year>) <article-title>Evolution of human growth spurts</article-title>. <source>Am J Phys Anthropol</source><volume>101</volume>: <fpage>455</fpage>–<lpage>474</lpage>.<pub-id pub-id-type="pmid">9016361</pub-id></mixed-citation></ref><ref id="pone.0107963-Smith1"><label>85</label><mixed-citation publication-type="journal"><name><surname>Smith</surname><given-names>RJ</given-names></name>, <name><surname>Jungers</surname><given-names>WL</given-names></name> (<year>1997</year>) <article-title>Body mass in comparative primatology</article-title>. <source>J Hum Evol</source><volume>32</volume>: <fpage>523</fpage>–<lpage>559</lpage>.<pub-id pub-id-type="pmid">9210017</pub-id></mixed-citation></ref><ref id="pone.0107963-Wood1"><label>86</label><mixed-citation publication-type="journal"><name><surname>Wood</surname><given-names>B</given-names></name> (<year>1976</year>) <article-title>The nature and basis of sexual dimorphism in the primate skeleton</article-title>. <source>J Zool</source><volume>180</volume>: <fpage>15</fpage>–<lpage>34</lpage>.</mixed-citation></ref><ref id="pone.0107963-Frayer1"><label>87</label><mixed-citation publication-type="journal"><name><surname>Frayer</surname><given-names>DW</given-names></name>, <name><surname>Wolpoff</surname><given-names>MH</given-names></name> (<year>1985</year>) <article-title>Sexual dimorphism</article-title>. <source>Annu Rev Anthropol</source><volume>14</volume>: <fpage>429</fpage>–<lpage>473</lpage>.</mixed-citation></ref><ref id="pone.0107963-Rogers1"><label>88</label><mixed-citation publication-type="book">Rogers J, Waldron T (1995) A field guide to joint disease in archaeology. Wiley, Chichester.</mixed-citation></ref><ref id="pone.0107963-MartinDupont1"><label>89</label><mixed-citation publication-type="journal"><name><surname>Martin-Dupont</surname><given-names>S</given-names></name>, <name><surname>Cunha</surname><given-names>E</given-names></name>, <name><surname>Rouge</surname><given-names>D</given-names></name>, <name><surname>Crubezy</surname><given-names>E</given-names></name> (<year>2006</year>) <article-title>Spondylarthropathy striking prevalence in a 19th–20th century Portuguese collection</article-title>. <source>Joint Bone Spine</source><volume>73</volume>: <fpage>303</fpage>–<lpage>310</lpage>.<pub-id pub-id-type="pmid">16564195</pub-id></mixed-citation></ref><ref id="pone.0107963-Lovell2"><label>90</label><mixed-citation publication-type="journal"><name><surname>Lovell</surname><given-names>NC</given-names></name> (<year>1997</year>) <article-title>Trauma analysis in paleopathology</article-title>. <source>Am J Phys Anthropol</source><volume>104</volume>: <fpage>139</fpage>–<lpage>170</lpage>.</mixed-citation></ref><ref id="pone.0107963-Ortner1"><label>91</label><mixed-citation publication-type="book">Ortner DJ, Putschar WGJ (1981) Identification of pathological conditions in human skeletal remains. Washington: Smithsonian Institution Press.</mixed-citation></ref><ref id="pone.0107963-Facchini1"><label>92</label><mixed-citation publication-type="other">Facchini F, Mariotti V, Bonfiglioli B, Belcastro MG (2006) Les collections ostéologiques et ostéoarchéologiques du musée d’Anthropologie de l’université de Bologne (Italie). Bulletin archéologique de Provence: 67–70.</mixed-citation></ref><ref id="pone.0107963-Bolter2"><label>93</label><mixed-citation publication-type="journal"><name><surname>Bolter</surname><given-names>DR</given-names></name>, <name><surname>Zihlman</surname><given-names>AL</given-names></name> (<year>2011</year>) <article-title>Brief communication: Dental development timing in captive Pan paniscus with comparisons to Pan troglodytes</article-title>. <source>Am J Phys Anthropol</source><volume>145</volume>: <fpage>647</fpage>–<lpage>652</lpage>.<pub-id pub-id-type="pmid">21541924</pub-id></mixed-citation></ref><ref id="pone.0107963-Molnar2"><label>94</label><mixed-citation publication-type="journal"><name><surname>Molnar</surname><given-names>S</given-names></name> (<year>1971</year>) <article-title>Human tooth wear, tooth function and cultural variability</article-title>. <source>Am J Phys Anthropol</source><volume>34</volume>: <fpage>175</fpage>–<lpage>189</lpage>.<pub-id pub-id-type="pmid">5572602</pub-id></mixed-citation></ref><ref id="pone.0107963-Henderson3"><label>95</label><mixed-citation publication-type="other">Henderson CY, Mariotti V, Pany-Kucera D, Perréard-Lopreno G, Villotte S, <etal>et al</etal> (2010) Scoring entheseal changes: proposal of a new standardised method for fibrocartilaginous entheses. Poster presented at the 18th European Meeting of the Paleopathology Association, Vienna, Austria 23rd–26th of August 2010. Consulted in 6th March 2013. Available: <ext-link ext-link-type="uri" xlink:href="https://www.uc.pt/en/cia/msm/Vienna2010.pdf">https://www.uc.pt/en/cia/msm/Vienna2010.pdf</ext-link>.</mixed-citation></ref><ref id="pone.0107963-Niinimki2"><label>96</label><mixed-citation publication-type="journal"><name><surname>Niinimäki</surname><given-names>S</given-names></name> (<year>2012</year>) <article-title>The relationship between musculoskeletal stress markers and biomechanical properties of the humeral diaphysis</article-title>. <source>Am J Phys Anthropol</source><volume>147</volume>: <fpage>618</fpage>–<lpage>628</lpage>.<pub-id pub-id-type="pmid">22331594</pub-id></mixed-citation></ref><ref id="pone.0107963-Milella3"><label>97</label><mixed-citation publication-type="book">Milella M (2010) Skeletal markers of activity: methodological and interpretative reflections after the study of the whole Frassetto Sassari identified skeletal collections. [Ph.D. Dissertation]. Bologna: University of Bologna.</mixed-citation></ref><ref id="pone.0107963-Milella4"><label>98</label><mixed-citation publication-type="other">Milella M, Zampetti S, Belcastro M, Mariotti V (2009) Carrying loads and making shoes: skeletal markers of activity in a sample of porters and shoemakers from contemporary Italy. In: Santos A, Alves Cardoso F, Assis S, Villotte S, editors. Workshop in musculoskeletal stress markers (MSM): limitations and anchievements in the reconstruction of past activity patterns: program-abstract book: Department of Anthropology University of Coimbra Centro de Investigação em Antropologia e Saúde. 31. Available: <ext-link ext-link-type="uri" xlink:href="https://estudogeral.sib.uc.pt/jspui/bitstream/10316/15351/1/Workshop%20in%20musculoskeletal%20stress%20markers.pdf">https://estudogeral.sib.uc.pt/jspui/bitstream/10316/15351/1/Workshop%20in%20musculoskeletal%20stress%20markers.pdf</ext-link>. Accessed 2013 Mar 31.</mixed-citation></ref><ref id="pone.0107963-Revelle1"><label>99</label><mixed-citation publication-type="other">Revelle W (2014) psych: Procedures for Personality and Psychological Research. Northwestern University, Evanston, Illinois, USA.</mixed-citation></ref><ref id="pone.0107963-R1"><label>100</label><mixed-citation publication-type="other">R Core Team (2012) R: A language and environment for statistical computing. Vienna, Austria.</mixed-citation></ref><ref id="pone.0107963-Leigh2"><label>101</label><mixed-citation publication-type="journal"><name><surname>Leigh</surname><given-names>SR</given-names></name>, <name><surname>Shea</surname><given-names>BT</given-names></name> (<year>1996</year>) <article-title>Ontogeny of body size variation in African apes</article-title>. <source>Am J Phys Anthropol</source><volume>99</volume>: <fpage>43</fpage>–<lpage>65</lpage>.<pub-id pub-id-type="pmid">8928723</pub-id></mixed-citation></ref><ref id="pone.0107963-Doran1"><label>102</label><mixed-citation publication-type="journal"><name><surname>Doran</surname><given-names>DM</given-names></name> (<year>1997</year>) <article-title>Ontogeny of locomotion in mountain gorillas and chimpanzees</article-title>. <source>J Hum Evol</source><volume>32</volume>: <fpage>323</fpage>–<lpage>344</lpage>.<pub-id pub-id-type="pmid">9085185</pub-id></mixed-citation></ref><ref id="pone.0107963-Rothschild1"><label>103</label><mixed-citation publication-type="journal"><name><surname>Rothschild</surname><given-names>BM</given-names></name>, <name><surname>Woods</surname><given-names>RJ</given-names></name> (<year>1989</year>) <article-title>Spondyloarthropathy in gorillas</article-title>. <volume>18</volume>: <fpage>267</fpage>–<lpage>276</lpage>.</mixed-citation></ref><ref id="pone.0107963-Jiang1"><label>104</label><mixed-citation publication-type="journal"><name><surname>Jiang</surname><given-names>Y</given-names></name>, <name><surname>Zhao</surname><given-names>J</given-names></name>, <name><surname>van Holsbeeck</surname><given-names>MT</given-names></name>, <name><surname>Flynn</surname><given-names>MJ</given-names></name>, <name><surname>Ouyang</surname><given-names>X</given-names></name>, <etal>et al</etal> (<year>2002</year>) <article-title>Trabecular microstructure and surface changes in the greater tuberosity in rotator cuff tears</article-title>. <source>Skeletal Radiol</source><volume>31</volume>: <fpage>522</fpage>–<lpage>528</lpage>.<pub-id pub-id-type="pmid">12195505</pub-id></mixed-citation></ref><ref id="pone.0107963-Milz1"><label>105</label><mixed-citation publication-type="journal"><name><surname>Milz</surname><given-names>S</given-names></name>, <name><surname>Tischer</surname><given-names>T</given-names></name>, <name><surname>Buettner</surname><given-names>A</given-names></name>, <name><surname>Schieker</surname><given-names>M</given-names></name>, <name><surname>Maier</surname><given-names>M</given-names></name>, <etal>et al</etal> (<year>2004</year>) <article-title>Molecular composition and pathology of entheses on the medial and lateral epicondyles of the humerus: a structural basis for epicondylitis</article-title>. <source>Ann Rheum Dis</source><volume>63</volume>: <fpage>1015</fpage>–<lpage>1021</lpage>.<pub-id pub-id-type="pmid">15308511</pub-id></mixed-citation></ref><ref id="pone.0107963-Hoyte1"><label>106</label><mixed-citation publication-type="journal"><name><surname>Hoyte</surname><given-names>DAN</given-names></name>, <name><surname>Enlow</surname><given-names>DH</given-names></name> (<year>1966</year>) <article-title>Wolff’s law and the problem of muscle attachment on resorptive surfaces of bone</article-title>. <source>Am J Phys Anthropol</source><volume>24</volume>: <fpage>205</fpage>–<lpage>213</lpage>.<pub-id pub-id-type="pmid">4957122</pub-id></mixed-citation></ref><ref id="pone.0107963-Drfl1"><label>107</label><mixed-citation publication-type="journal"><name><surname>Dörfl</surname><given-names>J</given-names></name> (<year>1980</year>) <article-title>Migration of tendinous insertions. I. Cause and mechanism</article-title>. <source>J Anat</source><volume>131</volume>: <fpage>179</fpage>.<pub-id pub-id-type="pmid">7440401</pub-id></mixed-citation></ref><ref id="pone.0107963-Drfl2"><label>108</label><mixed-citation publication-type="journal"><name><surname>Dörfl</surname><given-names>J</given-names></name> (<year>1980</year>) <article-title>Migration of tendinous insertions. II. Experimental modifications</article-title>. <source>J Anat</source><volume>131</volume>: <fpage>229</fpage>.<pub-id pub-id-type="pmid">7462092</pub-id></mixed-citation></ref><ref id="pone.0107963-Benjamin4"><label>109</label><mixed-citation publication-type="journal"><name><surname>Benjamin</surname><given-names>M</given-names></name>, <name><surname>Toumi</surname><given-names>H</given-names></name>, <name><surname>Suzuki</surname><given-names>D</given-names></name>, <name><surname>Redman</surname><given-names>S</given-names></name>, <name><surname>Emery</surname><given-names>P</given-names></name>, <etal>et al</etal> (<year>2007</year>) <article-title>Microdamage and altered vascularity at the enthesis–bone interface provides an anatomic explanation for bone involvement in the HLA–B27–associated spondylarthritides and allied disorders</article-title>. <source>Arthritis Rheum</source><volume>56</volume>: <fpage>224</fpage>–<lpage>233</lpage>.<pub-id pub-id-type="pmid">17195226</pub-id></mixed-citation></ref><ref id="pone.0107963-Benjamin5"><label>110</label><mixed-citation publication-type="journal"><name><surname>Benjamin</surname><given-names>M</given-names></name>, <name><surname>Milz</surname><given-names>S</given-names></name>, <name><surname>Bydder</surname><given-names>GM</given-names></name> (<year>2008</year>) <article-title>Magnetic resonance imaging of entheses. Part 1</article-title>. <source>Clin Radiol</source><volume>63</volume>: <fpage>691</fpage>–<lpage>703</lpage>.<pub-id pub-id-type="pmid">18455562</pub-id></mixed-citation></ref><ref id="pone.0107963-Lovejoy1"><label>111</label><mixed-citation publication-type="other">Lovejoy CO, Kern KF, Simpson SW, Meindl RS (1989) A new method for estimation of skeletal dimorphism in fossil samples with an application to Australopithecus afarensis. Proceedings of the 2nd International Congress of Human Paleoanthropology: 103–108.</mixed-citation></ref><ref id="pone.0107963-Richmond1"><label>112</label><mixed-citation publication-type="journal"><name><surname>Richmond</surname><given-names>BG</given-names></name>, <name><surname>Jungers</surname><given-names>WL</given-names></name> (<year>1995</year>) <article-title>Size variation and sexual dimorphism in Australopithecus afarensis and living hominoids</article-title>. <source>J Hum Evol</source><volume>29</volume>: <fpage>229</fpage>–<lpage>245</lpage>.</mixed-citation></ref><ref id="pone.0107963-Niinimki3"><label>113</label><mixed-citation publication-type="journal"><name><surname>Niinimäki</surname><given-names>S</given-names></name> (<year>2011</year>) <article-title>What do muscle marker ruggedness scores actually tell us?</article-title><source>Int J Osteoarch</source><volume>21</volume>: <fpage>292</fpage>–<lpage>299</lpage>.</mixed-citation></ref><ref id="pone.0107963-Peterson1"><label>114</label><mixed-citation publication-type="journal"><name><surname>Peterson</surname><given-names>J</given-names></name> (<year>1998</year>) <article-title>The Natufian hunting conundrum: spears, atlatls, or bows? Musculoskeletal and armature evidence</article-title>. <source>Int J Osteoarch</source><volume>8</volume>: <fpage>378</fpage>–<lpage>389</lpage>.</mixed-citation></ref></ref-list></back></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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