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