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Experimental evidence indicates that the lobes integrate visual and vestibular information and control the vestibulo-ocular reflex, vestibulo-collic reflex, smooth pursuit and gaze holding. Multiple paleoneuroanatomy studies have deduced the behaviour of fossil vertebrates by measuring the floccular fossae (FF). These studies assumed that there are correlations between FF volume and behaviour. However, these assumptions have not been fully tested. Here, we used micro-CT scans of extant mammals (47 species) and birds (59 species) to test six possible morphological-functional associations between FF volume and ecological\/behavioural traits of extant animals. Behaviour and ecology do not explain FF volume variability in four out of six variables tested. Two variables with significant results require further empirical testing. Cerebellum plasticity may explain the lack of statistical evidence for the hypotheses tested. Therefore, variation in FF volume seems to be better explained by a combination of factors such as anatomical and phylogenetic evolutionary constraints, and further empirical testing is required.<\/jats:p>","DOI":"10.1038\/s41598-017-01981-0","type":"journal-article","created":{"date-parts":[[2017,5,10]],"date-time":"2017-05-10T16:15:05Z","timestamp":1494432905000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":56,"title":["Floccular fossa size is not a reliable proxy of ecology and behaviour in vertebrates"],"prefix":"10.1038","volume":"7","author":[{"given":"S.","family":"Ferreira-Cardoso","sequence":"first","affiliation":[]},{"given":"R.","family":"Ara\u00fajo","sequence":"additional","affiliation":[]},{"given":"N. E.","family":"Martins","sequence":"additional","affiliation":[]},{"given":"G. G.","family":"Martins","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7002-1005","authenticated-orcid":false,"given":"S.","family":"Walsh","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1912-150X","authenticated-orcid":false,"given":"R. M. S.","family":"Martins","sequence":"additional","affiliation":[]},{"given":"N.","family":"Kardjilov","sequence":"additional","affiliation":[]},{"given":"I.","family":"Manke","sequence":"additional","affiliation":[]},{"given":"A.","family":"Hilger","sequence":"additional","affiliation":[]},{"given":"R.","family":"Castanhinha","sequence":"additional","affiliation":[]}],"member":"297","published-online":{"date-parts":[[2017,5,17]]},"reference":[{"key":"1981_CR1","doi-asserted-by":"publisher","first-page":"950","DOI":"10.1038\/nature02048","volume":"425","author":"LM Witmer","year":"2003","unstructured":"Witmer, L. M., Chatterjee, S., Franzosa, J. & Rowe, T. Neuroanatomy of flying reptiles and implications for flight, posture and behaviour. Nature \n                           425, 950\u2013953, doi:10.1038\/nature02048 (2003).","journal-title":"Nature"},{"key":"1981_CR2","doi-asserted-by":"publisher","first-page":"e80974","DOI":"10.1371\/journal.pone.0080974","volume":"8","author":"R Castanhinha","year":"2013","unstructured":"Castanhinha, R. et al. Bringing Dicynodonts Back to Life: Paleobiology and Anatomy of a New Emydopoid Genus from the Upper Permian of Mozambique. PLOS ONE \n                           8, e80974, doi:10.1371\/journal.pone.0080974 (2013).","journal-title":"PLOS ONE"},{"key":"1981_CR3","doi-asserted-by":"crossref","unstructured":"Balanoff, A. M. et al. Best practices for digitally constructing endocranial casts: examples from birds and their dinosaurian relatives. J. Anat. 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(University of Texas at Austin, 2004)."},{"key":"1981_CR7","doi-asserted-by":"publisher","first-page":"661","DOI":"10.1111\/j.1469-7580.2011.01427.x","volume":"219","author":"T Miyashita","year":"2011","unstructured":"Miyashita, T., Arbour, V. M., Witmer, L. M. & Currie, P. J. The internal cranial morphology of an armoured dinosaur Euoplocephalus corroborated by X-ray computed tomographic reconstruction. Journal of Anatomy \n                           219, 661\u2013675, doi:10.1111\/j.1469-7580.2011.01427.x (2011).","journal-title":"Journal of Anatomy"},{"key":"1981_CR8","doi-asserted-by":"publisher","first-page":"e67176","DOI":"10.1371\/journal.pone.0067176","volume":"8","author":"SA Walsh","year":"2013","unstructured":"Walsh, S. A. et al. Avian Cerebellar Floccular Fossa Size Is Not a Proxy for Flying Ability in Birds. PLOS ONE \n                           8, e67176, doi:10.1371\/journal.pone.0067176 (2013).","journal-title":"PLOS ONE"},{"key":"1981_CR9","first-page":"1","volume":"18","author":"DA Thomas","year":"2015","unstructured":"Thomas, D. A. The cranial anatomy of Tenontosaurus tilletti Ostrom, 1970 (Dinosauria, Ornithopoda). Palaeontol Electron \n                           18, 1\u201399 (2015).","journal-title":"Palaeontol Electron"},{"key":"1981_CR10","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1130\/SPE55-p1","volume":"55","author":"EC Olson","year":"1944","unstructured":"Olson, E. C. Origin of Mammals Based Upon Cranial Morphology of the Therapsid Suborders. Geological Society of America Special Papers \n                           55, 1\u2013130, doi:10.1130\/SPE55 (1944).","journal-title":"Geological Society of America Special Papers"},{"key":"1981_CR11","doi-asserted-by":"publisher","first-page":"1089","DOI":"10.1002\/jmor.20397","volume":"276","author":"M Laa\u00df","year":"2015","unstructured":"Laa\u00df, M. Virtual reconstruction and description of the cranial endocast of Pristerodon mackayi (Therapsida, Anomodontia). Journal of Morphology \n                           276, 1089\u20131099, doi:10.1002\/jmor.v276.9 (2015).","journal-title":"Journal of Morphology"},{"key":"1981_CR12","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1002\/ajpa.1330770202","volume":"77","author":"PJ Gannon","year":"1988","unstructured":"Gannon, P. J., Eden, A. R. & Laitman, J. T. The subarcuate fossa and cerebellum of extant primates: Comparative study of a skull-brain interface. Am. J. Phys. Anthropol. \n                           77, 143\u2013164, doi:10.1002\/ajpa.1330770202 (1988).","journal-title":"Am. J. Phys. Anthropol."},{"key":"1981_CR13","unstructured":"Larsell, O. & Jansen, J. The Comparative Anatomy and Histology of the Cerebellum, from Myxinoids Through Birds: By Olof Larsell. Ed. by Jan Jansen. (University of Minnesota Press, 1967)."},{"key":"1981_CR14","doi-asserted-by":"publisher","first-page":"275","DOI":"10.1146\/annurev.ne.05.030182.001423","volume":"5","author":"M Ito","year":"1982","unstructured":"Ito, M. Cerebellar Control of the Vestibulo-Ocular Reflex\u2013Around the Flocculus Hypothesis. Annual Review of Neuroscience \n                           5, 275\u2013297, doi:10.1146\/annurev.ne.05.030182.001423 (1982).","journal-title":"Annual Review of Neuroscience"},{"key":"1981_CR15","doi-asserted-by":"crossref","first-page":"1222","DOI":"10.1152\/jn.1994.71.3.1222","volume":"71","author":"DE Angelaki","year":"1994","unstructured":"Angelaki, D. E. & Hess, B. J. Inertial representation of angular motion in the vestibular system of rhesus monkeys. I. Vestibuloocular reflex. Journal of Neurophysiology \n                           71, 1222\u20131249 (1994).","journal-title":"Journal of Neurophysiology"},{"key":"1981_CR16","doi-asserted-by":"crossref","unstructured":"Voogd, J. & Barmack, N. H. In Progress in Brain Research (ed. B\u00fcttner-Ennever, J. A.) 151, 231\u2013268 (Elsevier, 2006).","DOI":"10.1016\/S0079-6123(05)51008-2"},{"key":"1981_CR17","first-page":"26","volume":"17","author":"NM Gerrits","year":"1989","unstructured":"Gerrits, N. M. & Voogd, J. The topographical organization of climbing and mossy fiber afferents in the flocculus and the ventral paraflocculus in rabbit, cat and monkey. Exp Brain Res Suppl \n                           17, 26\u201329 (1989).","journal-title":"Exp Brain Res Suppl"},{"key":"1981_CR18","doi-asserted-by":"crossref","first-page":"2945","DOI":"10.1152\/jn.2000.84.6.2945","volume":"84","author":"M Kahlon","year":"2000","unstructured":"Kahlon, M. & Lisberger, S. G. Changes in the Responses of Purkinje Cells in the Floccular Complex of Monkeys After Motor Learning in Smooth Pursuit Eye Movements. Journal of Neurophysiology \n                           84, 2945\u20132960 (2000).","journal-title":"Journal of Neurophysiology"},{"key":"1981_CR19","doi-asserted-by":"crossref","unstructured":"Voogd, J., Gerrits, N. M. & Hess, D. T. In Cerebellum and Neuronal Plasticity (eds Glickstein, M., Yeo, C. & Stein, J.) 15\u201340 (Springer US, 1987).","DOI":"10.1007\/978-1-4613-0965-9_2"},{"key":"1981_CR20","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1007\/BF00238229","volume":"50","author":"W Waespe","year":"1983","unstructured":"Waespe, W., Cohen, B. & Raphan, T. Role of the flocculus and paraflocculus in optokinetic nystagmus and visual-vestibular interactions: Effects of lesions. Exp Brain Res \n                           50, 9\u201333, doi:10.1007\/BF00238229 (1983).","journal-title":"Exp Brain Res"},{"key":"1981_CR21","doi-asserted-by":"crossref","first-page":"912","DOI":"10.1152\/jn.00768.2000","volume":"87","author":"H Rambold","year":"2002","unstructured":"Rambold, H., Churchland, A., Selig, Y., Jasmin, L. & Lisberger, S. G. Partial Ablations of the Flocculus and Ventral Paraflocculus in Monkeys Cause Linked Deficits in Smooth Pursuit Eye Movements and Adaptive Modification of the VOR. Journal of Neurophysiology \n                           87, 912\u2013924 (2002).","journal-title":"Journal of Neurophysiology"},{"key":"1981_CR22","doi-asserted-by":"publisher","first-page":"581","DOI":"10.1146\/annurev.neuro.27.070203.144238","volume":"27","author":"ES Boyden","year":"2004","unstructured":"Boyden, E. S., Katoh, A. & Raymond, J. L. Cerebellum-dependent learning: the role of multiple plasticity mechanisms. Annual Review of Neuroscience \n                           27, 581\u2013609, doi:10.1146\/annurev.neuro.27.070203.144238 (2004).","journal-title":"Annual Review of Neuroscience"},{"key":"1981_CR23","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3389\/fneur.2011.00053","volume":"2","author":"A Kheradmand","year":"2011","unstructured":"Kheradmand, A. & Zee, D. S. Cerebellum and ocular motor control. 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Journal of Vestibular Research \n                           17, 227\u2013237 (2007).","journal-title":"Journal of Vestibular Research"},{"key":"1981_CR26","unstructured":"Hegde, H. Eye Movements in Elite Athletes - An Index for Performance. Theses and Dissertations (2010)."},{"key":"1981_CR27","doi-asserted-by":"crossref","first-page":"1129","DOI":"10.1152\/jn.1986.56.4.1129","volume":"56","author":"H Noda","year":"1986","unstructured":"Noda, H. & Mikami, A. Discharges of neurons in the dorsal paraflocculus of monkeys during eye movements and visual stimulation. J. Neurophysiol. \n                           56, 1129\u20131146 (1986).","journal-title":"J. Neurophysiol."},{"key":"1981_CR28","doi-asserted-by":"publisher","first-page":"209","DOI":"10.1007\/s00221-006-0661-x","volume":"177","author":"I Kralj-Hans","year":"2007","unstructured":"Kralj-Hans, I., Baizer, J. S., Swales, C. & Glickstein, M. Independent roles for the dorsal paraflocculus and vermal lobule VII of the cerebellum in visuomotor coordination. Exp Brain Res \n                           177, 209\u2013222, doi:10.1007\/s00221-006-0661-x (2007).","journal-title":"Exp Brain Res"},{"key":"1981_CR29","doi-asserted-by":"publisher","first-page":"267","DOI":"10.1007\/s00359-007-0304-0","volume":"194","author":"AN Iwaniuk","year":"2007","unstructured":"Iwaniuk, A. N., Heesy, C. P., Hall, M. I. & Wylie, D. R. W. Relative Wulst volume is correlated with orbit orientation and binocular visual field in birds. J Comp Physiol A \n                           194, 267\u2013282, doi:10.1007\/s00359-007-0304-0 (2007).","journal-title":"J Comp Physiol A"},{"key":"1981_CR30","doi-asserted-by":"crossref","unstructured":"Jerison, H. J. Evolution of the Brain and Intelligence. (Academic Press, 1973).","DOI":"10.1016\/B978-0-12-385250-2.50018-3"},{"key":"1981_CR31","unstructured":"Hopson, J. A. Paleoneurology: pp. 39\u2013146 in C. Gans (ed.), Biology of the Reptilia Volume 9: Neurology A. (Academic Press, London, 1979)."},{"key":"1981_CR32","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1139\/z01-204","volume":"80","author":"AN Iwaniuk","year":"2002","unstructured":"Iwaniuk, A. N. & Nelson, J. E. Can endocranial volume be used as an estimate of brain size in birds? Can. J. Zool. \n                           80, 16\u201323, doi:10.1139\/z01-204 (2002).","journal-title":"Can. J. Zool."},{"key":"1981_CR33","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1671\/0272-4634(2007)27[99:DCEOPA]2.0.CO;2","volume":"27","author":"TE Macrini","year":"2007","unstructured":"Macrini, T. E., Muizon, C. D., Cifelli, R. L. & Rowe, T. Digital cranial endocast of Pucadelphys andinus, a Paleocene metatherian. 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Endocranial Morphology of the Primitive Nodosaurid Dinosaur Pawpawsaurus campbelli from the Early Cretaceous of North America. PLOS ONE \n                           11, e0150845, doi:10.1371\/journal.pone.0150845 (2016).","journal-title":"PLOS ONE"},{"key":"1981_CR41","unstructured":"Trotteyn, M. J. et al. Nuevas evidencias del desarrollo del fl\u00f3culo en el cerebelo de arcosauriformes: variabilidad e implicancias paleobiol\u00f3gicas. in ResearchGate (2015)."},{"key":"1981_CR42","unstructured":"Chatterjee, D. S. The Rise of Birds: 225 Million Years of Evolution. (The Johns Hopkins University Press, 1997)."},{"key":"1981_CR43","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1038\/nature12424","volume":"501","author":"AM Balanoff","year":"2013","unstructured":"Balanoff, A. M., Bever, G. S., Rowe, T. B. & Norell, M. A. Evolutionary origins of the avian brain. 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