{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,10,12]],"date-time":"2025-10-12T02:19:14Z","timestamp":1760235554820,"version":"build-2065373602"},"reference-count":47,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2021,9,7]],"date-time":"2021-09-07T00:00:00Z","timestamp":1630972800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100010665","name":"H2020 Marie Sk\u0142odowska-Curie Actions","doi-asserted-by":"publisher","award":["H2020-MSCA-IF-2018 No. 835571"],"award-info":[{"award-number":["H2020-MSCA-IF-2018 No. 835571"]}],"id":[{"id":"10.13039\/100010665","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Symmetry"],"abstract":"<jats:p>The observation and the quantification of asymmetry in biological structures are deeply investigated in geometric morphometrics. Patterns of asymmetry were explored in both living and fossil species. In living organisms, levels of directional and fluctuating asymmetry are informative about developmental processes and health status of the individuals. Paleontologists are primarily interested in asymmetric features introduced by the taphonomic process, as they may significantly alter the original shape of the biological remains, hampering the interpretation of morphological features which may have profound evolutionary significance. Here, we provide a new R tool that produces the numerical quantification of fluctuating and directional asymmetry and charts asymmetry directly on the specimens under study, allowing the visual inspection of the asymmetry pattern. We tested this show.asymmetry algorithm, written in the R language, on fossil and living cranial remains of the genus Homo. show.asymmetry proved successful in discriminating levels of asymmetry among sexes in Homo sapiens, to tell apart fossil from living Homo skulls, to map effectively taphonomic distortion directly on the fossil skulls, and to provide evidence that digital restoration obliterates natural asymmetry to unnaturally low levels.<\/jats:p>","DOI":"10.3390\/sym13091644","type":"journal-article","created":{"date-parts":[[2021,9,8]],"date-time":"2021-09-08T02:41:07Z","timestamp":1631068867000},"page":"1644","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":5,"title":["A New Integrated Tool to Calculate and Map Bilateral Asymmetry on Three-Dimensional Digital Models"],"prefix":"10.3390","volume":"13","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7919-4431","authenticated-orcid":false,"given":"Marina","family":"Melchionna","sequence":"first","affiliation":[{"name":"Dipartimento di Scienze della Terra, dell\u2019Ambiente e delle Risorse, Universit\u00e0 di Napoli Federico II, 80126 Napoli, Italy"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2884-7118","authenticated-orcid":false,"given":"Antonio","family":"Profico","sequence":"additional","affiliation":[{"name":"PalaeoHub, Department of Archaeology, Hull York Medical School University of York, Heslington YO10 5DD, UK"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8951-2990","authenticated-orcid":false,"given":"Costantino","family":"Buzi","sequence":"additional","affiliation":[{"name":"DFG Centre for Advanced Studies \u2018Words, Bones, Genes, Tools\u2019, Eberhard Karls University of T\u00fcbingen, 72070 T\u00fcbingen, Germany"}]},{"given":"Silvia","family":"Castiglione","sequence":"additional","affiliation":[{"name":"Dipartimento di Scienze della Terra, dell\u2019Ambiente e delle Risorse, Universit\u00e0 di Napoli Federico II, 80126 Napoli, Italy"}]},{"given":"Alessandro","family":"Mondanaro","sequence":"additional","affiliation":[{"name":"Dipartimento di Scienze della Terra, Universit\u00e0 degli Studi di Firenze, 50121 Firenze, Italy"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6620-4514","authenticated-orcid":false,"given":"Antonietta","family":"Del Bove","sequence":"additional","affiliation":[{"name":"Departament d\u2019Hist\u00f2ria i Hist\u00f2ria de l\u2019Art, Universitat Rovira i Virgili, 43002 Tarragona, Spain"},{"name":"Institut Catal\u00e0 de Paleoecologia Humana i Evoluci\u00f3 Social (IPHES-CERCA), 43007 Tarragona, Spain"}]},{"given":"Gabriele","family":"Sansalone","sequence":"additional","affiliation":[{"name":"Function, Evolution & Anatomy Research Lab, Zoology Division, School of Environmental and Rural Science, University of New England, Armidale, NSW 2351, Australia"}]},{"given":"Paolo","family":"Piras","sequence":"additional","affiliation":[{"name":"Dipartimento di Scienze Cardiovascolari, Respiratorie, Nefrologiche, Anestesiologiche e Geriatriche, Universit\u00e0 degli Studi di Roma La Sapienza, 00185 Rome, Italy"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4593-8006","authenticated-orcid":false,"given":"Pasquale","family":"Raia","sequence":"additional","affiliation":[{"name":"Dipartimento di Scienze della Terra, dell\u2019Ambiente e delle Risorse, Universit\u00e0 di Napoli Federico II, 80126 Napoli, Italy"}]}],"member":"1968","published-online":{"date-parts":[[2021,9,7]]},"reference":[{"key":"ref_1","first-page":"14","article-title":"A developmental perspective on developmental instability: Theory, models and mechanisms","volume":"Volume 1","author":"Polak","year":"2003","journal-title":"Developmental Instability: Causes and Consequences"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1046\/j.1095-8312.2003.00206.x","article-title":"Fluctuating asymmetry in bank vole populations (Rodentia, Arvicolinae) reflects stress caused by landscape fragmentation in the Mont-Saint-Michel Bay","volume":"80","author":"Marchand","year":"2003","journal-title":"Biol. J. Linn. Soc."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.jenvrad.2003.07.001","article-title":"High levels of fluctuating asymmetry in populations of Apodemus flavicollis from the most contaminated areas in Chornobyl","volume":"73","author":"Oleksyk","year":"2004","journal-title":"J. Environ. Radioact."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1086\/679011","article-title":"Effects of environmental disturbance on phenotypic variation: An integrated assessment of canalization, developmental stability, modularity, and allometry in lizard head shape","volume":"185","author":"Carretero","year":"2015","journal-title":"Am. Nat."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"524","DOI":"10.1016\/j.marpolbul.2014.09.038","article-title":"Marine pollution effects on the southern surf crab Ovalipes trimaculatus (Crustacea: Brachyura: Polybiidae) in Patagonia Argentina","volume":"91","author":"Lezcano","year":"2015","journal-title":"Mar. Pollut. Bull."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1007\/s10682-020-10051-3","article-title":"Fluctuating asymmetry in Alpine chamois horns: An indicator of environmental stress","volume":"34","author":"Chirichella","year":"2020","journal-title":"Evol. Ecol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"380","DOI":"10.1016\/j.evolhumbehav.2011.03.002","article-title":"Human fluctuating asymmetry in relation to health and quality: A meta-analysis","volume":"32","author":"Gangestad","year":"2011","journal-title":"Evol. Hum. Behav."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1002\/ajhb.22390","article-title":"Assessing the relationship between fluctuating asymmetry and cause of death in skeletal remains: A test of the developmental origins of health and disease hypothesis","volume":"25","author":"Weisensee","year":"2013","journal-title":"Am. J. Hum. Biol."},{"key":"ref_9","first-page":"187","article-title":"Female appearance: Facial and bodily attractiveness as shape","volume":"48","author":"Schaefer","year":"2006","journal-title":"Psychol. Sci."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"708","DOI":"10.1016\/j.evolhumbehav.2012.05.005","article-title":"Cues to fertility: Perceived attractiveness and facial shape predict reproductive success","volume":"33","author":"Oberzaucher","year":"2012","journal-title":"Evol. Hum. Behav."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1016\/j.evolhumbehav.2013.11.005","article-title":"Who is the fairest of them all? The independent effect of attractive features and self-perceived attractiveness on cooperation among women","volume":"35","author":"Pita","year":"2014","journal-title":"Evol. Hum. Behav."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"39731","DOI":"10.1038\/srep39731","article-title":"Predictors of facial attractiveness and health in humans","volume":"7","author":"Foo","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1016\/j.actpsy.2009.03.008","article-title":"Averageness or symmetry: Which is more important for facial attractiveness?","volume":"131","author":"Komori","year":"2009","journal-title":"Acta Psychol."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Graham, J.H., and \u00d6zener, B. (2016). Fluctuating asymmetry of human populations: A review. Symmetry, 8.","DOI":"10.3390\/sym8120154"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1002\/ajpa.22210","article-title":"Skull shape asymmetry and the socioeconomic structure of an early medieval central european society","volume":"150","author":"Bigoni","year":"2013","journal-title":"Am. J. Phys. Anthropol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"828","DOI":"10.1126\/science.1103707","article-title":"Symmetry breaking and the evolution of development","volume":"306","author":"Palmer","year":"2004","journal-title":"Science"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"505","DOI":"10.1038\/35107134","article-title":"Asymmetric Broca\u2019s area in great apes","volume":"414","author":"Cantalupo","year":"2001","journal-title":"Nature"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1111\/j.1469-7580.2012.01528.x","article-title":"Sexual dimorphism in multiple aspects of 3D facial symmetry and asymmetry defined by spatially dense geometric morphometrics","volume":"221","author":"Claes","year":"2012","journal-title":"J. Anat."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Ekrami, O., Claes, P., White, J.D., Weinberg, S.M., Marazita, M.L., Walsh, S., Shriver, M.D., and Van Dongen, S. (2020). A multivariate approach to determine the dimensionality of human facial asymmetry. Symmetry, 12.","DOI":"10.3390\/sym12030348"},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Palmer, A.R. (2005). Antisymmetry. Variation, Elsevier.","DOI":"10.1016\/B978-012088777-4\/50018-1"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Vingerhoets, G., Gerrits, R., and Verhelst, H. (2021). Atypical brain asymmetry in human situs inversus: Gut feeling or real evidence?. Symmetry, 13.","DOI":"10.3390\/sym13040695"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"2545","DOI":"10.1038\/srep02545","article-title":"Fossilization causes organisms to appear erroneously primitive by distorting evolutionary trees","volume":"3","author":"Sansom","year":"2013","journal-title":"Sci. Rep."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"160342","DOI":"10.1098\/rsos.160342","article-title":"Reconstructing the past: Methods and techniques for the digital restoration of fossils","volume":"3","author":"Lautenschlager","year":"2016","journal-title":"R. Soc. Open Sci."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1016\/j.jhevol.2009.04.004","article-title":"Principles for the virtual reconstruction of hominin crania","volume":"57","author":"Gunz","year":"2009","journal-title":"J. Hum. Evol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1016\/j.jhevol.2011.02.005","article-title":"A new OH5 reconstruction with an assessment of its uncertainty","volume":"61","author":"Benazzi","year":"2011","journal-title":"J. Hum. Evol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"154","DOI":"10.1002\/ajpa.22376","article-title":"Technical Note: Virtual reconstruction of KNM-ER 1813 Homo habilis cranium","volume":"153","author":"Benazzi","year":"2014","journal-title":"Am. J. Phys. Anthropol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1038\/nature03397","article-title":"Virtual cranial reconstruction of Sahelanthropus tchadensis","volume":"434","author":"Zollikofer","year":"2005","journal-title":"Nature"},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Tallman, M., Amenta, N., Delson, E., Frost, S.R., Ghosh, D., Klukkert, Z.S., Morrow, A., and Sawyer, G.J. (2014). Evaluation of a new method of fossil retrodeformation by algorithmic symmetrization: Crania of papionins (primates, cercopithecidae) as a test case. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0100833"},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Schlager, S., Profico, A., Di Vincenzo, F., and Manzi, G. (2018). Retrodeformation of fossil specimens based on 3D bilateral semi-landmarks: Implementation in the R package \u201cMorpho\u201d. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0194073"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"247","DOI":"10.3389\/feart.2020.00247","article-title":"Target Deformation of the Equus stenonis Holotype Skull: A Virtual Reconstruction","volume":"8","author":"Cirilli","year":"2020","journal-title":"Front. Earth Sci."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1093\/biomet\/87.2.285","article-title":"Statistical assessment of bilateral symmetry of shapes","volume":"87","author":"Mardia","year":"2000","journal-title":"Biometrika"},{"key":"ref_32","unstructured":"Zheng, G., Li, S., and Sz\u00e9kely, G. (2017). Morpho and Rvcg\u2014Shape Analysis in R: R-Packages for Geometric Morphometrics, Shape Analysis and Surface Manipulations. Statistical Shape and Deformation Analysis: Methods, Implementation and Applications, Academic Press."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"843","DOI":"10.3390\/sym7020843","article-title":"Analyzing fluctuating asymmetry with geometric morphometrics: Concepts, methods, and applications","volume":"7","author":"Klingenberg","year":"2015","journal-title":"Symmetry"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"eaax9935","DOI":"10.1126\/sciadv.aax9935","article-title":"Evolution of brain lateralization: A shared hominid pattern of endocranial asymmetry is much more variable in humans than in great apes","volume":"6","author":"Neubauer","year":"2020","journal-title":"Sci. Adv."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1002\/ajpa.24340","article-title":"Arothron: An R package for geometric morphometric methods and virtual anthropology applications","volume":"176","author":"Profico","year":"2021","journal-title":"Am. J. Phys. Anthropol."},{"key":"ref_36","unstructured":"Profico, A., Zeppilli, C., Micarelli, I., Mondanaro, A., Raia, P., Marchi, D., Manzi, G., and O\u2019Higgins, P. Morphometric maps of bilateral asymmetry in the human humerus: An implementation in the R package morphomap. Symmetry, under review."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"691","DOI":"10.1006\/jhev.1997.0163","article-title":"A comparative study of stereolithographically modelled skulls of Petralona and Broken Hill: Implications for future studies of middle Pleistocene hominid evolution","volume":"33","author":"Seidler","year":"1997","journal-title":"J. Hum. Evol."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Rightmire, G.P. (2017). Middle Pleistocene Homo Crania from Broken Hill and Petralona: Morphology, metric comparisons, and evolutionary relationships. Human Paleontology and Prehistory, Springer.","DOI":"10.1007\/978-3-319-46646-0_11"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"e1093","DOI":"10.7717\/peerj.1093","article-title":"Retrodeformation andmuscular reconstruction of ornithomimosaurian dinosaur crania","volume":"3","author":"Cuff","year":"2015","journal-title":"PeerJ"},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Buzi, C., Profico, A., Di Vincenzo, F., Harvati, K., Melchionna, M., Raia, P., and Manzi, G. (2021). Retrodeformation of the Steinheim cranium: Insights into the evolution of Neanderthals. Symmetry, 34.","DOI":"10.3390\/sym13091611"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"16022","DOI":"10.1073\/pnas.0904119106","article-title":"The origin of Neandertals","volume":"106","author":"Hublin","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1007\/s13219-015-0127-4","article-title":"The phylogenetic status of Homo heidelbergensis\u2014A cladistic study of Middle Pleistocene hominins","volume":"27","author":"Mounier","year":"2015","journal-title":"Bull. Mem. Soc. Anthropol. Paris"},{"key":"ref_43","first-page":"749","article-title":"Amira: A highly interactive system for visual data analysis","volume":"1","author":"Stalling","year":"2005","journal-title":"Vis. Handb."},{"key":"ref_44","first-page":"132","article-title":"Electronic removal of encrustations inside the steinheim cranium reveals paranasal sinus features and deformations, and provides a revised endocranial volume estimate","volume":"273","author":"Prossinger","year":"2003","journal-title":"Anat. Rec. Part B New Anat."},{"key":"ref_45","unstructured":"Adams, D., Collyer, M., Kaliontzopoulou, A., and Baken, E. (2021, July 21). Geomorph: Software for Geometric Morphometric Analyses. Available online: https:\/\/cran.r-project.org\/package=geomorph."},{"key":"ref_46","unstructured":"Schwartz, J.H., and Tattersall, I. (2005). The Human Fossil Record, John Wiley & Sons."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"643","DOI":"10.1002\/ar.a.20205","article-title":"CT-based description and phyletic evaluation of the archaic human calvarium from Ceprano, Italy","volume":"285","author":"Bruner","year":"2005","journal-title":"Anat. Rec. Part A Discov. Mol. Cell. Evol. Biol."}],"container-title":["Symmetry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-8994\/13\/9\/1644\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T06:57:47Z","timestamp":1760165867000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-8994\/13\/9\/1644"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,9,7]]},"references-count":47,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2021,9]]}},"alternative-id":["sym13091644"],"URL":"https:\/\/doi.org\/10.3390\/sym13091644","relation":{},"ISSN":["2073-8994"],"issn-type":[{"type":"electronic","value":"2073-8994"}],"subject":[],"published":{"date-parts":[[2021,9,7]]}}}