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.