DOI: 10.5061/dryad.v6wwpzh17
title: Conserved patterns and locomotor-related evolutionary constraints in the hominoid vertebral column

description: The evolution of the hominoid lineage is characterized by pervasive
 homoplasy, notably in regions such as the vertebral column, which plays a
 central role in body support and locomotion. Few isolated and fewer
 associated vertebrae are known for most fossil hominoid taxa, but
 identified specimens indicate potentially high levels of convergence in
 terms of both form and number. Homoplasy thus complicates attempts to
 identify the anatomy of the last common ancestor of hominins and other
 taxa and stymies reconstructions of evolutionary scenarios. One way to
 clarify the role of homoplasy is by investigating constraints via
 phenotypic integration, which assesses covariation among traits, shapes
 evolutionary pathways, and itself evolves in response to selection. We
 assessed phenotypic integration and evolvability across the subaxial
 (cervical, thoracic, lumbar, sacral) vertebral column of macaques (n =
 96), gibbons (n = 77), chimpanzees (n = 92), and modern humans (n = 151).
 We found a mid-cervical cluster that may have shifted cranially in
 hominoids, a persistent thoracic cluster that is most marked in
 chimpanzees, and an expanded lumbosacral cluster in hominoids that is most
 expanded in gibbons. Our results highlight the highly conserved nature of
 the vertebral column. Taxa appear to exploit existing patterns of
 integration and ontogenetic processes to shift, expand, or reduce cluster
 boundaries. Gibbons appear to be the most highly derived taxon in our
 sample, possibly in response to their highly specialized locomotion.
description: This dataset includes linear data from the subaxial vertebral
 column (cervical, thoracic, lumbar, and sacral) of 96 <em>Macaca
 mulatta</em>, 76 <em>Hylobates lar</em>, 92
 <em>Pan troglodytes</em>, and 29 <em>Homo
 sapiens</em> individuals.  Data were collected
 using a Mitutoyo Digital Caliper accurate to 0.02 mm and are reported in
 millimeters (mm). "." is used to denote a decimal throughout.
 Raw linear data are provided for all available elements for each
 individual. Missing data is noted with 'NA' and indicates the
 element or trait was missing. Intraobserver error ranges between
 1.06-1.28%.  Not all individuals display the modal
 formula for that taxon. Vertebral IDs have been adjusted as described in
 the associated manuscript and README file. Human data
 from the British Museum are anonymized.
description: Files are uploaded as .CSV files.
description: # Conserved patterns and locomotor-related evolutionary constraints in the
 hominoid vertebral column Catalina I. Villamil, Emily R. Middleton We
 include four separate files, one for each taxon: homo.csv, pan.csv,
 hylobates.csv, macaca.csv Data are provided in CSV files with each row
 representing one individual. Data are organized by column. All taxa have 7
 cervical elements modally. The modal formula for *Homo* is 12 thoracic, 5
 lumbar, and 5 sacral (12T:5L:5S) elements, for *Pan* is 13T:4L:6S, for
 *Hylobates* is 13T:5L:4S, and for *Macaca* is 12T:7L:3S. Vertebral IDs for
 individuals with nonmodal formulae have been adjusted as follows: * We
 removed T7 from analyses for individuals with an extra thoracic segment,
 and moved subsequent thoracic vertebrae up in the sequence (T8 treated as
 T7, T9 treated as T8, etc.) * We treated T7 as missing for individuals
 with fewer than the modal number of thoracic elements and moved subsequent
 elements down in nthe sequeence (T7 treated as T8, T8 treated as T9, etc.)
 * We removed L3 (in *Hylobates*) or L4 (in *Macaca*) for individuals with
 an extra lumbar segment and moved subsequent vertebrae up in the sequence
 (L5 treated as L4, etc.). * We treated L2 as missing in individuals with
 fewer than modal lumbar segments and moved subsequent elements down in the
 sequence (L3 treated as L4, etc.). This occurred only in Pan. * No
 adjustments were made to variation in sacral element number. Human data
 from the British Museum are anonymized. Human data from the Galloway
 Collection (Makerere University, College of Health Sciences) and Dart
 Collection (University of Witwatersrand), which were included in the
 associated manuscript, are excluded to align with those institution's
 policies. Please refer to the associated manuscript for more detailed
 descriptions of each measurement. Each measurement name is listed with the
 element name/number first, as described below: C+NUMBER Cervical +
 position (1 through 7, e.g. C3) T+NUMBER Thoracic + position (1 through 12
 or 13, e.g. T9) L+NUMBER Lumbar + position (1 through 4, 5, or 7, e.g. L3)
 S Sacral (whole sacrum) S1 First sacral element In the following key,
 'vert' stands for each element name/number as described above.
 Measurement key: * vert_SPINE Spine length, measured cranially from the
 margin of the laminae to the tip of the spine with the calipers parallel
 to the spine * vert_PREZ Pre-zygapophyseal (cranial articular facets)
 breadth, measured from the two most lateral points of the pre-zygapophyses
 * vert_POSTZ Post-zygapophyseal (caudal articular facets) breadth,
 measured from the two most lateral points of the post-zygapophyses *
 vert_PREZPOSTZ Interzygous height, measured from most cranial point of the
 pre-zygapophysis to the most caudal point of the post-zygapophysis *
 vert_BODY_HEIGHT Ventral craniocaudal body height, measured at midline *
 vert_BODY_BREADTH Caudal transverse body breadth, measured at the greatest
 breadth * vert_BODY_WIDTH Caudal dorsoventral body width, measured at the
 midline * vert_MAX_AUB Maximum transverse breadth at ventral projections
 of the auricular surfaces * vert_CAUD_AU Transverse breadth at the caudal
 margin of the auricular surface * vert_VENTL Ventral length, measured from
 the midpoint of the promotory of S1 to the midpoint of the most
 caudal/distal sacral element * vert_CCH Craniocaudal height of first
 sacral element, measured at the ventral midline ANONID indicates the
 anonymized ID number for that individual (humans/Homo) MUSID indicates the
 museum catalogue number (all other taxa) Institutions: CPRCMUS Caribbean
 Primate Research Center Museum, San Juan, PR, USA MCZ Museum of
 Comparative Zoology, Harvard University, Cambridge, MA, USA AMNH American
 Museum of Natural History, New York, NY, USA HTB Hamann-Todd Osteological
 Collection, Cleveland Museum of Natural History, Cleveland, OH, USA
 Powell-Cotton Powell-Cotton Museum, Birchington-on-Sea, England, UK RG
 Royal Museum of Central Africa, Tervuren, Belgium USNM Smithsonian Museum
 of Natural History (US National Museums), Washington DC, USA Sex: F Female
 M Male U Unknown