TABLE 1. Genera and species represented, repository information, sample sizes, and specimen numbers used in this study. U Wits: University of the Witwatersrand, Johannesburg, South Africa; UCMP: University of California Museum of Paleontology, Berkeley, California, USA; Ditsong: Ditsong National Museum of Natural History, Pretoria, South Africa. TM211 was originally described as Papio spelaeus (Broom, 1936). M3147 has also been published as P. h. robinsoni (Gilbert et al., 2015).
| Genus | Species | Repository | N | Specimen numbers | Site(s) | References |
| Cercopithecoides | C. molletti | Ditsong | 1 | SK551 | Swartkrans | McKee et al. (2011) |
| C. williamsi | Ditsong | 3 | KA195, KB5241C, MP36 | Kromdraii, Makapansgat | Szalay and Delson (1979), McKee et al. (2011), Geissler (2013) | |
| U Wits | 1 | STS394A | Sterkfontein | Geissler (2013) | ||
| Total | 5 | |||||
| Dinopithecus | D. ingens | Ditsong | 5 | SK553, SK554, SK599, SK600, SK604 | Swartkrans | Gilbert (2013) |
| Total | 5 | |||||
| Gorgopithecus | G. major | Ditsong | 3 | KA153, KA605, KA944 | Kromdraii | Gilbert (2013) |
| Total | 3 | |||||
| Papio | P. angusticeps | Ditsong | 5 | CO100, CO101, CO102, CO135A, KA194 | Cooper's Cave, Kromdraii | McKee and Keyser (1994), Williams et al. (2007), Gilbert (2013) |
| P. h. robinsoni | Ditsong | 5 | SK555, SK557, SK558, SK560, SK562 | Swartkrans | Carter (2006), Geissler (2013), Beaudet et al. (2016) | |
| UCMP | 2 | UCMP 56767, UCMP 56797 | Bolt's Farm | Freedman (1965) | ||
| U Wits | 1 | SWP31 | Sterkfontein | Eisenhart (1974) | ||
| P. h. ursinus | Ditsong | 1 | TM211 | Freedman (1957) | ||
| P. izodi | U Wits | 5 | T89-11-1, TP4, TP7, TP10, TP12 | Taung | Gilbert and Grine (2010), Gilbert (2013) | |
| Total | 19 | |||||
| Parapapio | P. broomi | Ditsong | 6 | STS254A, STS262, STS393, STS396A, STS397, STS564 | Sterkfontein | Carter (2006), Williams et al. (2007), Gilbert (2013) |
| UCMP | 1 | UCMP 56786 | Bolt's Farm | Freedman (1965) | ||
| U Wits | 7 | M3056, M3065, MP151, MP2, MP224, MP76, SWP12 | Makapansgat, Sterkfontein | Fourie (2006), Fourie et al. (2008), Gilbert (2013) | ||
| P. jonesi | Ditsong | 2 | STS260, STS565 | Sterkfontein | Pickering et al. (2004), Fourie et al. (2008), Gilbert (2013) | |
| U Wits | 5 | M3060, MP75, MP173, SWP389, SWP2947 | Makapansgat, Sterkfontein | Maier (1970) | ||
| P. whitei | UCMP | 1 | UCMP 56614 | Buxton Limeworks | Freedman (1965) | |
| U Wits | 10 | BF43, M3070, M3072, M3147, MP119, MP164, MP208, MP221, MP223, MP239 | Bolt's Farm, Makapansgat | Maier (1977), Williams et al. (2007), Fourie et al. (2008), Williams (2013) | ||
| unknown | U Wits | 4 | M3053, M3084, M3133, MP47 | Makapansgat | Fourie (2006), Fourie et al. (2008) | |
| Total | 36 | |||||
| Procercocebus | P. antiquus | UCMP | 3 | UCMP 56609, UCMP 56694, UCMP 125956 | Buxton Limeworks | Gilbert (2007) |
| U Wits | 5 | M3078, M3079, T89-154, TP8, TP9 | Makapansgat, Taung | Gilbert (2013) | ||
| Total | 8 | |||||
| Theropithecus | T. darti | U Wits | 4 | M3073, MP168, MP217, MP222 | Makapansgat | Maier (1972), Williams et al. (2007), Geissler (2013), Gilbert (2013) |
| T. oswaldi | Ditsong | 2 | SK561, SK563 | Swartkrans | Folinsbee and Reisz (2013) | |
| Total | 6 | |||||
| unknown | unknown | Ditsong | 6 | SB2, SK3, SK556, SK814, SKII25, STS537 | Swartkrans, Sterkfontein | N/A |
| UCMP | 1 | UCMP 218738 | Buxton Limeworks | Monson et al. (2015) | ||
| U Wits | 10 | unnumbered black skull, MP4A, SP1, SP24, SWP1728, SWP1731, SWP29, SWP2953, T8817, T9087 | Makapansgat, Sterkfontein, Taung | N/A | ||
| Total | 17 | |||||
| TOTAL | 99 |
TABLE 2. Extant papionin species represented and sample sizes used in this study. All specimens are from the National Museum of Natural History in Washington D.C., USA, with the exception of one Theropithecus specimen from the National Museum in Addis Ababa, Ethiopia.
| Species | Sample size (n) |
| Cercocebus torquatus | 20 |
| Lophocebus albigena | 15 |
| Mandrillus sphinx | 7 |
| Papio hamadryas | 34 |
| Theropithecus gelada | 5 |
| Total | 81 |
TABLE 3. Definitions of cranial measurements.
| Measurement | Definition |
| Maximum length | Maximum length of the cranium from prosthion to inion. |
| Maximum breadth | Maximum breadth of the cranium posterior to the external auditory meatus. |
| Orbital width | Maximum width of the orbit along a line perpendicular to the orbital height. |
| Orbital height | Maximum height of the orbit along a parasagittal plane. |
| Interorbital breadth | Breadth between the superomedial margins of the orbits. |
| Nasal width | Maximum width of the nasal aperture, from left to right alare. |
| Nasal height | Maximum height of the nasal aperture, from nasospinale to rhinion. |
| Muzzle width (ectomolare) | Maximum width of the muzzle, measured at the ectomolare. |
| Muzzle width (maxillary fossae) | Maximum width of the muzzle, measured at the superior border of the maxillary fossae, inferior to the orbits. |
| Maximum width (canine) | Maximum width of the muzzle, measured at the canine alveoli. |
| Maximum zygofrontal width | Maximum width of the upper face, measured from left to right frontomalare temporale. |
| Maximum calvarial width (post-orbital constriction) | Maximum calvarial width, measured at the post-orbital constriction. |
| Calvarial length | Maximum calvarial length, measured from ophryon to inion. |
| Palate width (M3) | Maximum width of the palate, measured from the lateral edges of the depth of the palate, along a line bisecting the third molars. |
| Palate width (canine) | Maximum width of the palate, measured from the lateral edges of the depth of the palate, along a line bisecting the most medial points of the canine alveoli. |
| Palate length | Maximum length of the palate, measured from orale to staphylion. |
| Face length | Projection of the face, measured from prosthion to glabella. |
TABLE 4. Sample size, mean, and standard deviation of each measurement for the fossil genera sampled in this study. All measurements are in millimeters.
| Genus |
Cercopithecoides |
Dinopithecus |
Gorgopithecus |
Papio |
Parapapio |
Procercocebus |
Theropithecus |
Unknown |
|||||||||
| Sample size | 5 | 5 | 3 | 19 | 36 | 8 | 6 | 17 | |||||||||
| Statistic | Mean | SD | Mean | SD | Mean | SD | Mean | SD | Mean | SD | Mean | SD | Mean | SD | Mean | SD | |
| Maximum length | 124.9 | NA | 173.9 | 21.7 | NA | NA | 148.2 | 9.8 | 152.4 | 19.9 | 147.9 | 16.5 | 171.5 | 8.1 | 163.9 | 30.6 | |
| Maximum breadth | 81.6 | 6.2 | 87.1 | 5.9 | NA | NA | 71.6 | 2.0 | 76.2 | 5.5 | 70.9 | 7.2 | 84.1 | 5.3 | 77.6 | 6.6 | |
| Orbital width | 29.4 | 2.2 | 30.1 | 6.0 | NA | NA | 27.5 | 1.9 | 26.9 | 2.2 | 26.4 | 1.6 | 28.7 | 2.4 | 27.0 | 2.3 | |
| Orbital height | 27.9 | 0.8 | 30.1 | 6.0 | NA | NA | 24.3 | 3.0 | 24.5 | 2.4 | 23.5 | 1.5 | 26.9 | 4.9 | 25.3 | 3.5 | |
| Interorbital | 15.0 | 2.9 | 11.9 | 5.9 | NA | NA | 11.5 | 2.7 | 11.4 | 2.0 | 8.9 | 1.2 | 12.1 | 1.5 | 11.8 | 2.4 | |
| Nasal width | 14.9 | 2.5 | 16.1 | 3.5 | 16.4 | NA | 19.1 | 4.0 | 17.6 | 2.4 | 17.7 | 1.8 | 18.9 | 1.7 | 18.0 | 2.0 | |
| Nasal height | 18.8 | NA | 27.6 | 1.8 | NA | NA | 29.8 | 9.1 | 26.5 | 4.2 | 25.8 | 2.7 | 29.3 | 2.4 | 29.3 | 8.0 | |
| Muzzle width (ectomolare) |
44.6 | 2.2 | 58.6 | 10.2 | 58.2 | NA | 51.4 | 8.0 | 51.2 | 4.2 | 46.8 | 2.4 | 51.9 | 3.1 | 48.1 | 6.6 | |
| Muzzle width (maxillary fossae) |
25.5 | 3.9 | 39.9 | 5.8 | NA | NA | 40.0 | 8.9 | 37.1 | 4.4 | 36.4 | 4.2 | 35.5 | 4.5 | 39.4 | 5.4 | |
| Maximum width (C) | 30.6 | 2.7 | 38.7 | 9.4 | 43.0 | 3.4 | 41.4 | 11.3 | 36.6 | 5.8 | 35.2 | 4.1 | 33.8 | 2.1 | 38.6 | 4.8 | |
| Maximum zygofrontal width | 81.0 | 3.2 | 88.1 | 21.3 | NA | NA | 72.4 | 6.5 | 72.4 | 6.9 | 75.8 | 4.9 | 77.1 | 1.1 | 80.5 | 5.4 | |
| Maximum cal. width | 54.9 | 3.8 | 54.1 | 0.9 | NA | NA | 53.4 | 5.8 | 53.3 | 3.0 | 56.4 | 3.6 | 48.2 | 4.7 | 53.4 | 3.5 | |
| Calvarial length | 94.4 | 10.7 | 92.3 | NA | NA | NA | 88.5 | 3.5 | 94.3 | 8.3 | 87.9 | 4.7 | 95.4 | 10.5 | 90.8 | 6.6 | |
| Palate width (M3) | 19.2 | 1.5 | 25.1 | 6.9 | 25.9 | 7.1 | 23.9 | 5.2 | 24.1 | 2.3 | 20.5 | 2.7 | 24.3 | 4.7 | 21.2 | 2.9 | |
| Palate width (C) | 18.8 | 1.8 | 27.8 | 8.1 | 23.6 | 0.0 | 24.2 | 6.2 | 21.5 | 3.0 | 17.8 | 2.8 | 20.4 | 1.1 | 22.0 | 3.0 | |
| Palate length | 40.4 | 2.1 | 68.9 | 10.5 | 65.5 | 4.4 | 67.4 | 16.9 | 51.5 | 8.2 | 51.2 | 6.0 | 62.9 | 3.8 | 57.6 | 11.7 | |
| Face length | 62.4 | 6.5 | 98.7 | 29.7 | NA | NA | 99.0 | 30.7 | 82.2 | 11.6 | 76.1 | 10.6 | 93.0 | 0.9 | 94.0 | 28.6 | |
TABLE 5. Results of the MANOVA for the extant and fossil papionins. Double asterisks denote significance at p < 0.001.
| Traits | Group | p |
| Max. length, Max. breadth, Orb. width, Orb. height, Interorb. breadth, Nas. width, Nas. height, Muz. width (ecto.), Muz. width (max. fos.), Max. cal. width | Extant | 0.000** |
| Max. length, Max. breadth, Orb. width, Orb. height, Interorb. breadth, Nas. width, Nas. height, Muz. width (ecto.), Muz. width (max. fos.), Max. cal. width | Fossil | 0.439 |
| Cal. length, Palate width (M3), Palate width (C), Palate length, Face length | Extant | 0.000** |
| Cal. length, Palate width (M3), Palate width (C), Palate length, Face length | Fossil | 0.382 |
TABLE 6. ANOVA results for the size-corrected values of the extant sample, n = 81. MANOVA is significant at p < 2.2e-16. Measurements are size-corrected using geometric mean (Mosimann, 1970). Shaded cells are significant at p < 0.05.
| Genus comparison | Muz. Width (Ecto.) |
Muz. Width (Canine) |
Palate Width (Canine) | Palate Length | Face Length |
| Lophocebus-Cercocebus | 0.087 | 0.017 | 0.017 | 0.980 | 0.036 |
| Mandrillus-Cercocebus | 0.000 | 0.712 | 0.712 | 0.000 | 0.002 |
| Papio-Cercocebus | 0.000 | 0.040 | 0.040 | 0.000 | 0.000 |
| Theropithecus-Cercocebus | 1.000 | 0.224 | 0.224 | 0.000 | 0.000 |
| Mandrillus-Lophocebus | 0.000 | 0.007 | 0.007 | 0.000 | 0.000 |
| Papio-Lophocebus | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 |
| Theropithecus-Lophocebus | 0.522 | 0.001 | 0.001 | 0.000 | 0.000 |
| Papio-Mandrillus | 0.192 | 0.987 | 0.987 | 0.991 | 0.024 |
| Theropithecus-Mandrillus | 0.000 | 0.938 | 0.938 | 0.007 | 0.438 |
| Theropithecus-Papio | 0.000 | 0.986 | 0.986 | 0.000 | 0.957 |
TABLE 7. ANOVA results for a subset of the size-corrected values of the fossil sample, n = 38. MANOVA is significant at p < 0.0009. Measurements are size-corrected using geometric mean (Mosimann, 1970). Shaded cells are significant at p < 0.05.
| Genus Comparison | Muz. Width (Ecto.) |
Muz. Width (Canine) |
Palate Width (Canine) | Palate Length | Face Length |
| Dinopithecus-Cercopithecoides | 0.404 | 0.687 | 0.991 | 0.140 | 0.355 |
| Papio-Cercopithecoides | 0.035 | 0.966 | 0.930 | 0.004 | 0.300 |
| Parapapio-Cercopithecoides | 0.876 | 0.998 | 1.000 | 0.980 | 0.853 |
| Procercocebus-Cercopithecoides | 0.855 | 0.999 | 0.504 | 0.301 | 0.771 |
| Theropithecus-Cercopithecoides | 0.733 | 0.152 | 0.647 | 0.011 | 0.088 |
| Papio-Dinopithecus | 0.975 | 0.895 | 1.000 | 0.966 | 0.998 |
| Parapapio-Dinopithecus | 0.682 | 0.651 | 0.981 | 0.103 | 0.642 |
| Procercocebus-Dinopithecus | 0.855 | 0.318 | 0.892 | 0.938 | 0.882 |
| Theropithecus-Dinopithecus | 0.958 | 0.963 | 0.954 | 0.982 | 0.998 |
| Parapapio-Papio | 0.005 | 0.979 | 0.607 | 0.000 | 0.421 |
| Procercocebus-Papio | 0.099 | 0.605 | 0.800 | 0.167 | 0.912 |
| Theropithecus-Papio | 0.292 | 0.150 | 0.932 | 1.000 | 0.842 |
| Procercocebus-Parapapio | 1.000 | 0.837 | 0.090 | 0.169 | 0.997 |
| Theropithecus-Parapapio | 0.983 | 0.025 | 0.236 | 0.001 | 0.089 |
| Theropithecus-Procercocebus | 0.999 | 0.011 | 1.000 | 0.342 | 0.400 |
TABLE 8. Sample size and correlations for a subset of size-corrected measurement comparisons. A full correlation matrix comparing all size-corrected measurements is available in Appendix 8. All correlations are significant at p < 0.0004. “(C)” is the abbreviation for canine.
| Measurement pair | Sample size (n) |
Correlation (r) |
| Maximum length - face length | 31 | 0.898 |
| Maximum length - palate length | 35 | 0.880 |
| Maximum zygofrontal width - orbital width | 32 | 0.835 |
| Palate length - face length | 45 | 0.827 |
| Maximum length - maximum breadth | 29 | 0.817 |
| Maximum length - calvarial length | 31 | 0.814 |
| Maximum width (C) - palate width (C) | 54 | 0.798 |
| Calvarial length - face length | 29 | 0.784 |
| Calvarial length - maximum breadth | 39 | 0.770 |
| Muzzle width ectomolare - palate width (C) | 54 | 0.764 |
| Face length - nasal height | 35 | 0.760 |
| Maximum width (C) - face length | 43 | 0.758 |
