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TABLE 1. All Homunculus specimensa recovered by the MLP-Duke expeditions, 2003 to 2019.

Year of Discovery Catalog Numberb Element Loc.c Collector Publications Figure in Publication
2003 MPM-PV 3504 Partial mandible with Li2, p2-3, Rm1-3 ELC Jonathan Perry (Perry et al., 2010; Kay et al., 2012) figs. 2, 8; figs. 16.10c, 16.11b, c
2004 MPM-PV 3500 Right humerus missing proximal end PLC Jonathan Perry (Kay et al., 2012; Perry et al., 2018; Fleagle et al., 2022) fig. 16.15; None; figs. 2-10
2004 MPM-PV 3502 Cranium missing left superior part of braincase and left zygomatic arch KAN Carlos Luna (Perry et al., 2010; Kay et al., 2012; This Publication) figs. 1, 2, 8; figs. 16.2c, 16.3b, 16.4a, 16.8a, b, d, 16.11a; Figure 8
2004 MPM-PV 3501 Cranium missing anterior part of palate; partly distorted PLC Jonathan Perry (Perry et al., 2010; Kay et al., 2012) figs. 1, 9; fig. 16.2a
2004 MPM-PV 3503 Left half of cranium with associated upper right cheek teeth PLC Richard Kay (Kay et al., 2012) fig. 16.2b
2007 MPM-PV 3505 Face, palate, and anterior cranial fossae of a subadult (with deciduous premolars) PLC Jonathan Perry (Kay et al., 2012; Perry et al., 2014) figs. 1, 4-10; figs. 16.2d-h
2010 MPM-PV 3708 Right hemimandible with p2-4, m1-3, and intact condyle PLC Juan Carlos Fernicola (Kay et al., 2012; Kay and Perry, 2019; Li et al., 2020) fig. 16.12b; fig. 3.3
2011 MPM-PV 4376 Left mandible with m1-m2 and anterior alveoli RBu Leonel Acosta and Sergio Vizcaíno (Li et al., 2020) None
2013 MPM-PV 19426* Left mandibular fragment with m1 BB Néstor Toledo (Kay and Perry, 2019; Li et al., 2020) figs. 5.1, 6
2014 MPM-PV 19427* Right mandible with m1-3 (all broken) SBB Jackson Spradley (Kay and Perry, 2019) None
2014 MPM-PV 19428* Left mandibular fragment with m1-2 SBB Laura Chornogubsky (Kay and Perry, 2019) fig. 5.2
2015 MPM-PV 17453 Partial skeleton with associated skull PLC Sergio Vizcaíno This Publication; Li et al., 2020 Figure 3, Figure 5, Figure 8, Figure 11, Figure 12, Figure 13, Figure 14
2017 MPM-PV 17452 Left m3 RBo Susana Bargo and Sergio Vizcaíno (Kay and Perry, 2019) figs. 3.4, 5.3

aFor a list of specimens discovered by others, see Kay et al. (2012).
bAll specimens are accessioned at the Museo Regional Provincial "Padre Manuel Jesús Molina" (Río Gallegos, Santa Cruz Province, Argentina). Specimens are listed in order of discovery.
cSee Figure 2 for map of localities. BB, Barrancas Blancas, Río Santa Cruz; ELC, Estancia La Costa; KAN, Killik Aike Norte; PLC, Puesto Estancia La Costa; RBo, Río Bote; RBu, Rincón del Buque; SBB, Segundas Barrancas Blancas, Río Santa Cruz.
Asterisk indicates specimens that belong to Homunculus vizcainoi; all others are H. patagonicus.

 

TABLE 2. Prediction variables for Homunculus chewing muscle PCSA and mass.

Soft Tissue Variable Osteological Proxy
Temporalis PCSA Temporalis Insertion Area, measured from surface of mandible (TIA)
Temporalis Origin Area, measured from surface of cranium (TOA)
Masseter PCSA Masseter Insertion Area, measured from surface of mandible (MIA)
Masseter Origin Area, measured from surface of cranium (MOA)
Medial Pterygoid PCSA Medial Pterygoid Insertion Area, measured from surface of mandible (MPIA)
Medial Pterygoid Origin Area, measured from surface of cranium (MPOA)
Temporalis Mass TIA x To-ia
Masseter Mass MIA x Mo-i
Medial Pterygoid Mass MPIA x MPo-i

aTo-i is the distance between the centroid of the area of origin to the centroid of the area of insertion for the temporalis. This is measured the same way for Mo-i (masseter) and MPo-i (medial pterygoid).

 

 

TABLE 3. Cranial measurementsa of select Homunculus specimens.

  MPM-PV 17453 MPM-PV 3501 MPM-PV 3502 MPM-PV 3503 MPM-PV 3505 MPM-PV 5000b
Cranial Length 75.58 68.82* 70.45* 72.87* n/a n/a
Cranial Breadth PBc 37.57 33.22 34.82 27.04* 31.81 30.16
Cranial Breadth PCd 31.03 24.39 25.28 28.28* 27.70 27.80
Facial Heighte 10.85 11.71 10.82 11.15 8.06 9.48
Orbit Heightf 14.03 14.85 14.66 16.60 12.75 14.01
Orbit Breadthg 14.38 14.30 13.45 9.78* 12.16 12.23
Interorbital Breadthh 6.11 5.06 4.20 5.52* 3.53 4.40
Orbital Convergencei 68 n/a 70 n/a n/a 64
Foramen Magnum Lengthj 11.56 11.38 9.82 n/a n/a n/a
Foramen Magnum Breadthk 9.59 6.81* 9.14 n/a n/a n/a
Palate Lengthl 28.29 25.46 28.44 23.36* 24.99 25.80
Palate Breadthm 22.53 22.96 24.23 23.12 23.24 22.74
Geometric Meann 18.43 16.88 17.00 18.85 14.76 15.60
Maxillary Deptho 7.45 7.93 10.93 n/a n/a 11.54
Pre-orbital Rostral Lengtho 15.34 16.37 15.7 n/a n/a 14.42

a All measurements are in millimeters except orbital convergence (degrees). Asterisk indicates the best estimate given distortion or breakage. This measurement is the maximum distance in the sagittal plane.
b Type of “Killikaike blakei”
c Maximum right-to-left measurement at the postorbital bars.
d Outside caliper measurement at deepest part of postorbital constriction.
e Vertical distance from inferior edge of orbital rim to inferior edge of alveolar process of maxilla at the distal (=posterior) edge of the P4.
f Maximum inside dimension in vertical plane.
g Maximum inside dimension in horizontal plane. Orbital measurements taken from the less distorted orbit of the two.
h Distance between the most medial point on the left orbit and the most medial point on the right orbit.
i In degrees, as measured by Heesy, Ross, and colleagues (e.g., Heesy, 2003; Ross, et al., 2007).
j Maximum inside dimension in the antero-posterior plane.
k Maximum inside dimension in the mediolateral plane.
l Distal edge of M3 to inferior nasal spine; direct distance.
m Buccal edge of RM1 to buccal edge of LM1.
n Geometric mean of all above available measurements. Missing measurements were not estimated. This underestimates the true size for the very incomplete specimens MPM-PV 3505 and MPM-PV 5000.
o See Ross (1994).

 

TABLE 4. Lower tooth dimensionsa for Homunculus patagonicus, including MPM-PV 17453.

Specimen number i1 m-d i1 m-d root i1 b-l i2 m-d i2 m-d root i2 b-l /c m-d /c b-l /c height
MPM-PV 17453 1.62 n/a 2.06 2.62 n/a 2.43 3.48 3.28 4.12
MPM-PV 3504 1.88 1.49 2.04 2.68 1.53 2.45 3.91 2.84 6.11
MACN-SC 3114 1.62   2.32            
MPM-PV 3708             3.54 3.6  
MACN-A 634b             3 3.1  
  p2 m-d p3 b-l p3 m-d p4 b-l p4 m-d p4 b-l      
MPM-PV 17453 4.62 2.76 3.34 3.4 3.6 3.7      
MPM-PV 3504 3.68 3.05              
MPM-PV 3708 3.02 3.36 3.47 3.15 3.56 3.54      
MACN-A 634b 3.5 3.2 3.2 2.7 3.2 3.1      
MACN-A 5757c 3.34 2.67 3.42 2.85 3.08 3.38      
MACN-SC 2916     3.2 4.78          
MACN-SC 3026         3.83 3.34      
MACN-SC 3116 3.51 2.98              
  m1 m-d m1 trigonid b-l m1 talonid b-l m2 m-d m2 tri b-l m2 tal b-l m3 m-d m3 tri b-l m3 tal b-l
MPM-PV 17453 4.37 4.13 4.34 4.45 3.94 3.82 4.24 3.82 3.14
MPM-PV 3504       5.12 4.67 4.69 4.28 3.78 2.94
MPM-PV 3708 4.81 4.1 4.19 4.84 3.96 3.75 4.73 3.87 3.3
MPM-PV 4376 4.18 3.58 3.51 4.24 3.39 3.31      
MLP-PV 11-121 4.23 3.52 3.67            
MLP-PV 55-XII-13-156 4.24 3.39 3.75            
MACN-SC 3200           3.48      
MACN-SC 334 4.55 3.55 3.58            
MACN-SC 336 4.28 3.51 3.76 4.35 3.5 3.45      
MACN-SC 338 4.46 3.62 3.41            
MACN-SC 339 4.3 3.69 4.03 4.29 3.72 3.61      
MACN-SC 341 4.41 3.58 3.74            
MACN-SC 3074 4.32                
MACN-SC 3090       4.27 3.4 3.36      
MACN-SC 2918       4.31 3.56 3.4      
MACN-SC 1149       4.28 3.46 3.34      
MPM-PV 19428* 4.12 3.5 3.51 4.31 3.5 3.51      
MPM-PV 17452             4.73 3.55 2.86
MACN-A 10403 4.42 3.59 4 4.32 3.69 3.65      
MACN-A 5757c 4.38 3.5 3.8 4.27 3.49 3.6 4.23 3.45 2.92
MACN-A 5966       4.24 3.63 3.45      
MACN-A 5969d 4.43 3.71 3.87 4.4 3.59 3.64      
MACN-A 634b 4 2.8 3.2 4.5 3 3.5      
MPM-PV 17487       4.88 3.7 3.51      
MPM-PV 19426* 4.17 3.35 3.44            

a All measurements are in millimeters.
b From Rusconi (1935)
c Right side.
d Composite of a and b.
Asterisk indicates specimens that belong to H. vizcainoi, including the Type specimen (MPM-PV 19426); all others are H. patagonicus.

 

TABLE 5. Upper tooth dimensionsa for Homunculus patagonicus, including MPM-PV 17453.

Specimen number I1 m-d I1 m-d root I1 b-l I2 m-d I2 m-d root I2 b-l C/ m-d C/ b-l C/ height
MPM PV 17453b                  
MPM-PV 3502 3.45 2.58 2.16 3.22 2.24 2.09 3.48 2.97 3.99
MACN-A 5968             3.61 3.22  
MACN-SC 275             3.87 3.36 4.76
MACN-SC 335 (or 355)             3.88 3.18 5.23
MACN-SC 3089 2.95 1.82 1.95            
                   
MPM PV 5000             3.34 2.685 4.98
  P2 m-d P3 b-l P3 m-d P4 b-l P4 m-d P4 b-l
MPM-PV 17453b 2.62 3.89 3 4.67 3.32 5.13
MPM-PV 3501 3.24 3.91 2.93 4.51 3.38 5.23
MPM-PV 3502 2.47 3.62 2.98 4.51 3.11 4.9
MPM-PV 3503 3.28 3.67 3.13 4.3 3.14 4.99
CORD-PZ 1130 2.7 3.5 2.6 4.4 2.9 5.1
MACN-SC 2916     3.17 4.76    
MACN-SC 3116 3.56 3.25        
MPM-PV 5000 2.795 3.46 2.98 3.87    
  M1 m-d M1 b-l M2 m-d M2 b-l M3 m-d M3 b-l
MPM-PV 17453b 4.58 5.95 4.2 5.88 3.02 4.87
MPM-PV 3501 4.27 6.2 4.15 6.06 3.03 5.04
MPM-PV 3502 4.19 6.28 4.15 5.91 2.96 4.77
MPM-PV 3503 4.59 6.15 3.91 6.07 2.94 4.77
CORD-PZ 1130 3.77   3.75   2.62  
MACN-SC 274         3.07 4.8
MACN-SC 334     3.98 5.34    
MACN-SC 337     3.83 5.17    
MACN-SC 342     3.78 5.42    
MACN-SC 3012     4.01      
MPM-PV 5000 4.29   3.80 4.84 2.87 4.38
MPM-PV 1607c 4.1 5.7 3.7 4.9 2.9 4.5

a All measurements are in millimeters.
b Mean of left and right values, except for I2 for which the left-side tooth is lost.
c Measurements from Tejedor et al. (2008).
 

TABLE 6. Measurementsa of humeri of Homunculus.

  MPM-PV
17453 (L)
MPM-PV
17453 (R)
MPM-PV
3500
Maximum length 82.07* 98.61 92
Maximum midshaft diameter 9.08 8.71 8.37
M-d midshaft diameter 6.32 6.42 6.64
A-p midshaft diameter 7.15 7.35 6.20
Bi-epicondylar width 21.49 20.43 20.24
Medial epicondyle angle 22 21 22
Capitulum depth 7.35 6.64 7.01
Humeral head superoinferior n/a 13.01 n/a
Humeral distal mediolateral 15.28 14.54 14.30

a Humeral head measurements are used to estimate body mass (see Perry et al., 2018). Except for the humeral midshaft diameters, all others are from Fleagle et al. (2022), based on measurements described in Fleagle and Simons (1982; 1995). All measurements in millimeters except medial epicondyle angle which is in degrees. Asterisk denotes a minimum measurement in the case of breakage.

 

TABLE 7. Measurements of the scapula of MPM-PV 17453.

Measurement Left Scapula Right Scapula
Glenoid fossa height 12.03mm 11.93mm
Glenoid fossa width 7.60mm 7.69mm
Axillo-glenoid angle 138° 135°
Axillo-spinal angle 32° 30°
Spino-glenoid angle 106° 105°

 

TABLE 8. Predicted values for Homunculus chewing muscle PCSA and mass.

Y var. (log) X var.
 (log)a
RMA
r-squared
LS
slope
LS
 intercept
LS
SEEb
LS
QMLEc
Homunculus
predictedd
Temporalis PCSA TIAd 0.278 0.425 0.549 0.277 1.039 4.287
TOAe 0.566 0.859 -0.393 0.215 1.023 2.649
Masseter PCSA MIAf 0.369 0.738 0.353 0.215 1.023 5.860
MOAg 0.459 0.940 1.064 0.327 1.055 3.345
MP PCSA MPIAh 0.057 0.208 -0.097 0.328 1.055 0.928
MPOAi 0.088 0.418 0.113 0.323 1.054 1.041
Temporalis Mass TIA x To-ij 0.438 0.453 0.452 0.298 1.046 4.931
Masseter Mass MIA x Mo-i 0.719 0.813 0.057 0.275 1.039 4.618
MPMass MPIA x MPo-i 0.235 0.383 -0.392 0.351 1.064 0.643

a See Table 2 for explanation of abbreviations.
b Standard Error of the Estimate (=Root Mean Square Error). See Perry et al. (2018) for explanation of prediction procedure.
c Quasi-Maximum Likelihood Estimator. See Perry et al. (2018) for explanation of prediction procedure.
d Predicted values for PCSA are in cm2 and those for muscle volume are in g.

 

TABLE 9. Dimensions of chewing muscles for extant platyrrhines and those estimated for Homunculus.

Species Log GM
(cm)a
Temporalis Masseter Medial Pterygoid
Log PCSA
(cm2)
Log Mass
(g)
Log PCSA
 (cm2)
Log Mass
 (g)
Log PCSA
(cm2)
Log Mass
(g)
Aotus nancymaae 0.506 0.230 0.721 0.297 0.247 -0.335 -0.674
Callithrix jacchus 0.354 0.230 0.278 0.059 -0.059 -0.121 -0.509
Callithrix pygmaea 0.264 -0.018 -1.012 0.166 -1.261 -0.111 -2.234
Chiropotes sagulatus 0.682 0.783 0.816 1.213 1.222 0.303 0.217
Mico argentatus 0.320 0.207 0.031 0.089 -0.234 -0.359 -0.757
Saguinus imperator 0.352 0.233 -0.409 0.199 0.079 -0.389 -0.620
Saguinus labiatus 0.387 0.366 0.350 0.178 0.097 -0.210 -0.347
Saguinus midas 0.398 -0.368 -0.353 -0.482 -0.740 -0.645 -1.081
Saguinus oedipus 0.365 0.096 0.358 -0.066 -0.252 -0.263 -0.538
Saimiri sciureus 0.443 0.318 0.420 0.066 0.009 -0.037 -0.222
Sapajus apella 0.627 0.833 0.494 0.621 0.513 0.478 0.222
Homunculus patagonicus 0.530 0.423 0.693 0.524 0.664 n/a n/a

a The geometric mean is composed of the following measurements: mandible length, cranial length, corpus height below m2, maximum skull height in articulation, max orbit height, max bizygomatic breadth, cranial breadth behind zygomatic arches, breadth at the postorbital constriction, max bi-orbital breadth.