TABLE 1. Measurements taken on distal humeri in this study. Columns (from left to right) are: measurements (abbreviation and name); measurement description; taxon of study; and source reference for each measurement. We took the following measurements (Figure 2) on all multituberculate and therian specimens, contingent upon specimen completeness (see Table 2). The anatomical view from which the measurement was taken is indicated in parentheses in the description, as it relates to either multituberculate (Mu) or therian (Th) humeri, see text for more details. Measurements are identical to those in the indicated sources, with the following exceptions: our measurement TDW is ‘BW’ of Szalay and Dagosto (1980) and measurement 5 of Argot (2001); NCW is measurement 6 of Argot (2001) and ‘EEC’ of Boyer et al. (2010); TLW is ‘TLE’ of Szalay and Dagosto (1980); measurements TD and CD are modified from Szalay and Dagosto (1980) and Boyer et al. (2010). Multituberculate measurements lacking references are based on the structurally (but not necessarily functional) analogous measurement for therian humeri (see Figure 2).

Measurements Description Taxon Source
TDW Total Distal Width Medial edge of specimen to lateral edge, measured at the maximum extensions of the entepicondyle and ectepicondyle; parallel to other distal width measures (anterior view in therians; ventral view in multituberculates) Th, Mu Deischl 1964; Szalay and Dagosto 1980; Argot 2001
AW Articular Surface Width Eu: Lateral edge of capitulum to medial edge of trochlea (anterior)

Mu: Lateral edge of radial condyle to medial edge of ulnar condyle (ventral)

Th, Mu Szalay and Dagosto 1980; Szalay and Sargis 2001
TLW Total Lateral Width Eu: Medial edge of trochlea to lateral edge of ectepicondyle (anterior)

Mu: Medial edge of ulnar condyle to lateral edge of ectepicondyle (ventral)

Th, Mu Szalay and Dagosto 1980
NCW Entepicondylar Width Eu: Medial edge of trochlear lip to medial edge of entepicondyle (anterior)

Mu: From midpoint of groove between entepicondyle and ulnar condyle to medial edge of entepicondyle (ventral)

Th, Mu Argot 2001; Boyer et al. 2010
TW Trochlear Width Medial edge of trochlea (trochlear lip) to lateral edge of trochlea (medial edge of intercondylar groove) (anterior) Th Szalay and Dagosto 1980; Boyer et al. 2010
CW Capitulum Width Lateral edge of capitulum to medial edge of intercondylar groove (anterior) Th Szalay and Dagosto 1980; Boyer et al. 2010
TL Trochlear Length Length from proximal margin to distal margin of trochlear lip (anterior) Th Szalay and Dagosto 1980; Boyer et al. 2010
CL Capitulum Length Length from proximal margin to distal margin of capitulum (anterior) Th  this study
TD Trochlear Depth Maximum length from anterior surface to posterior surface of trochlea, parallel to CD (distal) Th Szalay and Dagosto 1980; Boyer et al. 2010
CD Capitulum Depth Length of capitulum articular surface, from anterior to posterior extent, parallel to TD (distal) Th Szalay and Dagosto 1980; Boyer et al. 2010
UCW Ulnar Condyle Width Medial edge of ulnar condyle to lateral edge of ulnar condyle (ventral) Mu  this study
RCW Radial Condyle Width Medial edge of radial condyle to lateral edge of radial condyle (ventral) Mu  this study
UCL Ulnar Condyle Length Length from proximal margin to distal margin of ulnar condyle (ventral) Mu  this study
RCL Radial Condyle Length Length from proximal margin to distal margin of radial condyle (ventral) Mu  this study
UCD Ulnar Condyle Depth Maximum length from ventral surface to dorsal surface ulnar condyle, parallel to RCD (distal) Mu  this study
RCD Radial Condyle Depth Maximum length from ventral surface to dorsal surface radial condyle, parallel to UCD (distal) Mu  this study

 

TABLE 2. Linear measurements of multituberculate distal humeri specimens. See Table 1 for measurement descriptions; all measurements on multituberculate humeri are taken in ventral view except for RCD and UCD, which are taken in distal view. All measurements are in millimeters. Abbreviations: Fm, Formation; HC, Hell Creek Formation; La, Lancian Biozone; Lan, Lance Formation; Morph., Morphotype; Pu1, Puercan 1 Biozone; Pu3, Puercan 3 Biozone; Tu, Tullock Formation. Asterisks indicate minimum size values, where preserved morphology was measured but the full measurement was not possible, due to specimen breakage; “-” indicates specimen was too broken for measurement; “?” indicates more tentative assignment to morphotype, due to specimen breakage. Locality numbers follow the system of each institution; those beginning with “V” are UCMP localities; those beginning with “C” are UWBM localities (see Appendix 1 for additional locality details).

Specimen Locality Fm Biozone Morph. TDW AW TDW NCW UCW RCW UCL RCL UCD RCD
195999 V5620 Lan La MuA 3.695 2.060 2.550 1.045 0.950 1.115 1.365 1.195 1.625 1.525
195974 V65127 Tu BCA MuA 3.995 2.005 2.455 1.255 0.765 1.000 1.305 1.140 1.530 1.510
195982 V65127 Tu BCA MuA - 2.040 2.210 - 0.865 1.170 1.405 1.055 1.445 -
122045 V70201 Tu BCA MuA 4.120 2.105 2.565 1.375 0.875 1.235 1.315 1.165 1.650 1.580
196956 V70201 Tu BCA MuA - 1.960* - - 0.780 1.120* 1.225 1.005 1.275 -
196957 V70201 Tu BCA MuA 3.915 2.090 2.615 1.175 0.840 1.160 1.465 1.240 1.460 1.455
70976 C0338 Tu BCA MuA - 1.850 2.420 - 0.855 1.070 1.345 1.150 1.440 1.400
70977 C0338 Tu BCA MuA 3.695 - 2.580 1.075 1.065 - 1.480 - 1.415 -
195998 V5620 Lan La ?MuA - - - - 0.690 - 0.805 - 1.805 -
153016 V74111 Tu Pu1 ?MuA - 2.165* - - - 1.240 1.475* 1.240* 1.585 1.475*
195933 V99438 Tu Pu3 MuB - - - 1.240 1.080 - 1.465 - 1.670 -
174490 V73087 HC La MuC - 3.555* - - 1.690 2.055 2.440 1.980 2.845 2.445
195997 V5620 Lan La MuC - 2.525 3.150 - 0.985 1.360 1.925 1.600 2.145 2.000
174420 V70209 Tu BCA MuC 5.290 2.975 3.595 1.480 1.295 1.630 1.975 2.030* 2.130 1.975
153039 V84193 HC Pu1 MuC 6.460 3.090 4.250 1.960 1.570 1.700 2.185 1.830 2.555 2.335
197687 V74111 Tu Pu1 MuC - 3.565* - - 1.480* 2.050* 2.240 1.885 2.330 -
195932 V99438 Tu Pu3 MuC 5.350* 3.035 3.820 1.460* 1.505 1.640 2.180 1.765 2.515 2.295
195945 V74122 Tu Pu3 MuC 5.590* 3.040 3.815 1.480 1.270 1.420 2.455 1.755 2.345 2.305
195990 V5620 Lan La MuD 7.310* 4.230* 5.470* 1.835* - 2.160 - 2.135 - 2.350
174439 V70209 Tu BCA MuE - - - 2.385 1.855 - 3.265 - 3.200 -
174442 V70209 Tu BCA MuE - - - - 1.965 - 2.730* - 2.600* 2.680
195973 V65127 Tu BCA MuE 8.130 4.025 5.080 2.445 1.485 2.410 2.810 2.265 3.200 3.145
97031 C1845 Tu BCA MuE 8.420 4.400 5.230 2.595 2.000 2.565 2.695 2.485 3.355 3.335
174412 V84162 HC Pu1 MuE - - - - 2.360 - 3.535 - 3.910 -
127384 V87001 HC La MuF 14.025 7.730 9.990 3.970 3.865 4.435 5.440 5.095 6.420 5.580
174400 V85092 HC La MuF - 6.900 - - 3.135 3.945 4.985 4.260 5.180 4.530
174232 V72207 HC La MuF - - - - - 5.050 - 5.655 - 6.365
174225 V72207 HC La MuF - 6.815* - 3.255 3.435 4.215* 5.045 4.900 5.835 -
174218 V73097 HC La MuF - 7.820* - - 2.450 4.325 5.605 5.145 6.420 -
101970 C1521 HC La MuF 13.705 6.945 9.105 3.995 2.985 3.915 5.030 3.900 5.030 4.405

 

 

TABLE 3. Linear measurements of therian distal humeri specimens. See Table 1 for measurement descriptions; all measurements on therian humeri are taken in anterior view except for CD and TD, which are taken in distal view . All measurements are in millimeters. Abbreviations: Fm, Formation; HC, Hell Creek Formation; La, Lancian Biozone; Morph., Morphotype; Pu1, Puercan 1 Biozone; Pu3, Puercan 3 Biozone; Tu, Tullock Formation. Asterisks indicate minimum size values, where preserved morphology was measured but the full measurement was not possible, due to specimen breakage; “-” indicates specimen was broken; “?” indicates more tentative assignment to morphotype; “‡” indicates specimen was used in 2D GM locomotor analyses.Locality numbers follow the system of each institution: those beginning with “V” are UCMP localities; those beginning with “C” are UWBM localities (see Appendix 1 for additional locality details). Five additional specimens from the BCA biozone (UCMP 151945, 151965, 151983, and 151984 from loc. V65127, and UCMP 174429 from loc. V70209) are tentatively attributed to EuD but are not listed below as they lack preservation sufficient for measurement, see text for more details.

Specimen Locality Fm Biozone Morph. TDW AW TLW NCW TW CW TL CL TD CD
127391 V88007 HC La ThA 4.900* - - 2.380 1.210 - 1.340 - 1.380 -
152394 V93169 HC La MeA 18.235* - - - - - - - - -
153100 V99438 Tu Pu3 EuA 6.400* 3.850 3.850 2.900* 1.330 2.406 1.630 1.579 1.756 1.920
97114‡ C1115 HC La EuB 8.53* 4.765 5.120 3.400* 1.705 2.895 1.960 1.945 2.355 2.335*
151991‡ V71203 HC BCA EuC 9.520 5.635 5.735 3.820 1.670 3.945 2.680 2.515 3.030 3.310
153023 V74111 Tu Pu1 EuC - - - 4.015 1.510* - 2.500* - 3.160 -
151985 V70201 Tu BCA EuD 7.810* 5.295 5.285 - 1.890 3.255 2.175 .720 2.235 2.495
151944 V65127 Tu BCA EuD 8.845* 5.735 6.005 2.190* 2.185 3.655 1.940 2.320 2.230* 2.575*
151946 V65127 Tu BCA EuD 8.800* 5.270 6.060 - 1.970 3.255 1.665 2.080 2.185 2.510*
151964‡ V65127 Tu BCA EuD 9.225 5.340 5.700 3.550 1.750 3.585 1.645 2.005 2.225 2.380
153036‡ V65127 Tu BCA EuD 9.935 5.675 5.850 4.210 1.820 3.800 1.775 2.155 2.470 2.710
218901 V91065 Tu Pu1 EuE 17.925* 10.685 10.850 7.175* 2.950 7.620 4.575 5.110 6.280 7.525
92928 C1163 Tu Pu3 EuE 15.030* 9.550 9.700 - 2.640 6.960 4.330 4.580 5.120* 5.320
153089 V72129 Tu Pu3 EuF 19.415* 17.280 17.395 - 5.480 11.820 6.945 7.280 8.735* 9.670
192678 V72124 Tu Pu3 EuG - - - 10.300 5.095 - 7.435 - 9.425* -

 

 

TABLE 4. Modern mammalian specimens used in geometric morphometrics analyses. Species list and locomotor references are modified from Chen and Wilson 2015; see text for more details. All specimens are from the UWBM except for those with specimen numbers A2001 and S2008, which are taken from images of specimens in distal view in Argot (2001, figure 8) and Salton and Sargis (2008, figure 4.10), respectively. Code number assigned (and table sorted) as follows: clade, order, family, and locomotor mode. Locomotor mode abbreviations are as follows: A, arboreal; F, fossorial; G, gliding; S, saltatorial; Sa, semiaquatic; Sc, scansorial; Sf, semifossorial; T, terrestrial. Other abbreviations: F, female; LM, locomotor mode; M, male; Spec., specimen number; U, sex unknown.

Clade Order Code LM Genus Species Family Spec. Sex Locomotor Reference
Metatheria
  Dasyuromorphia
    1 T Antechinus swainsonii Dasyuridae 68900 M Nowak 1999
    2 T Sarcophilus harrisii Dasyuridae 20671 M Van Valkenburgh 1987
  Didelphimorphia
    3 A Caluromys philander Didelphidae A2001 U Argot 2001; Nowak 1999
    4 A Caluromys derbianus Didelphidae 32255 U Argot 2001
    5 A Micoureus demerarae Didelphidae A2001 U Argot 2001
    6 Sc Didelphis virginiana Didelphidae 35525 F Argot 2001
    7 Sc Didelphis virginiana Didelphidae 74661 M Argot 2001
    8 Sc Didelphis marsupialis Didelphidae A2001 U Argot 2001; Nowak 1999
    9 Sc Marmosa murina Didelphidae A2001 U Argot 2001; Nowak 1999
    10 T Metachirus nudicaudatus Didelphidae 35438 F Argot 2001; Delciellos and Vieira 2006
    11 T Monodelphis brevicaudata Didelphidae A2001 U Argot 2001; Nowak 1999
  Diprotodontia
    12 G Petaurus breviceps Petauridae 34181 M Smith 1973; Nowak 1999; Körtner and Geiser 2000
    13 G Petaurus breviceps Petauridae 72928 U Smith 1973; Nowak 1999; Körtner and Geiser 2000
    14 A Trichosurus vulpecula Phalangeridae 68913 M Nowak 1999
    15 S Potorous tridactylus Potoroidae 41025 F Bassarova et al. 2009
    16 S Potorous tridactylus Potoroidae 34200 M Bassarova et al. 2009
    17 S Aepyprymnus rufescens Potoroidae 68898 F Bassarova et al. 2009
    18 A Pseudocheirus peregrinus Pseudocheiridae 68924 F Bassarova et al. 2009
    19 A Pseudocheirus peregrinus Pseudocheiridae 68908 M Bassarova et al. 2009
  Peramelemorphia
    20 T Perameles nasuta Peramelidae 68919 F Nowak 1999
Eutheria
  Afrosoricida
    21 Sa Potamogale velox Potamogalidae S2008 U Nowak 1999; Salton and Sargis 2008
    22 A Dendrohyrax arboreus Procaviidae 39039 F Nowak 1999
    23 A Dendrohyrax arboreus Procaviidae 39038 M Nowak 1999
    24 Sa Limnogale mergulus Tenrecidae S2008 U Nowak 1999; Salton and Sargis 2008
    25 Sc Echinops telfairi Tenrecidae 34168 F Endo et al. 2006; Salton and Sargis 2008
    26 Sc Echinops telfairi Tenrecidae S2008 U Endo et al. 2006; Salton and Sargis 2008
    27 Sf Hemicentetes semispinosus Tenrecidae S2008 U Nowak 1999; Salton and Sargis 2008
    28 Sf Oryzorictes sp. Tenrecidae S2008 U Nowak 1999; Salton and Sargis 2008
    29 T Setifer setosus Tenrecidae S2008 U Salton and Sargis 2008
    30 T Tenrec ecaudatus Tenrecidae S2008 U Salton and Sargis 2008
    31 T Microgale cowani Tenrecidae S2008 U Nowak 1999; Salton and Sargis 2008
    32 T Microgale dobsoni Tenrecidae S2008 U Nowak 1999; Salton and Sargis 2008
    33 T Microgale talazaci Tenrecidae S2008 U Nowak 1999; Salton and Sargis 2008
  Carnivora
    34 T Vulpes vulpes Canidae 39490 F Lariviére and Pasitschniak-Arts 1996; Meachen-Samuels 2010
    35 T Vulpes vulpes Canidae 39489 M Lariviére and Pasitschniak-Arts 1996; Meachen-Samuels 2010
    36 Sc Urocyon cinereoargenteus Felidae 77676 F Van Valkenburgh 1987; Meachen-Samuels 2010
    37 Sc Urocyon cinereoargenteus Felidae 35221 M Van Valkenburgh 1987; Meachen-Samuels 2010
    38 Sc Suricata suricatta Herpestidae 35470 F van Staaden 1994; Iwaniuk et al. 1999
    39 Sc Suricata suricatta Herpestidae 35469 M van Staaden 1994; Iwaniuk et al. 1999
    40 Sf Mephitis mephitis Mephitidae 35951 F Wade-Smith and Verts 1982; Van Valkenburgh 1987; Samuels and Van Valkenburgh 2008
    41 Sf Mephitis mephitis Mephitidae 39321 M Wade-Smith and Verts 1982; Van Valkenburgh 1987; Samuels and Van Valkenburgh 2008
    42 Sa Lontra canadensis Mustelidae 32226 F Gingerich 2003
    43 Sa Lontra canadensis Mustelidae 32230 M Gingerich 2003
    44 Sa Mustela vison Mustelidae 41780 F Howell 1930; Larivié́re 1999; Gingerich 2003
    45 Sa Mustela vison Mustelidae 35223 M Howell 1930; Larivié́re 1999; Gingerich 2003
    46 Sc Gulo gulo Mustelidae 14200 F Van Valkenburgh 1987; Pasitschniak-Arts and Larivié́re 1995
    47 Sc Gulo gulo Mustelidae 34417 M Van Valkenburgh 1987; Pasitschniak-Arts and Lariviére 1995
    48 Sf Spilogale putorius Mustelidae 39331 F Heinrich and Houde 2006
    49 Sf Spilogale putorius Mustelidae 39155 M Heinrich and Houde 2006
    50 T Mustela erminea Mustelidae 39366 F Nowak 1999
    51 T Mustela erminea Mustelidae 72863 M Nowak 1999
    52 T Mustela putorius Mustelidae 32599 F Nowak 1999
    53 T Mustela putorius Mustelidae 58727 M Nowak 1999
    54 A Paguma larvata Viverridae 73281 U Nowak 1999
  Cingulata
    55 F Cabassous centralis Dasypodidae 34167 F Nowak 1999
  56 F Dasypus novemcinctus Dasypodidae 20735 F Samuels and Van Valkenburgh 2008
  57 F Dasypus novemcinctus Dasypodidae 22458 M Samuels and Van Valkenburgh 2008
  58 F Euphractus sexcinctus Dasypodidae 35468 M Redford and Wetzel 1985; Nowak 1999
  Eulipotyphla
    59 T Echinosorex gymnurus Erinaceidae S2008 U Nowak 1999; Salton and Sargis 2008
  60 Sf Solenodon paradoxus Solenodontidae S2008 U Nowak 1999; Salton and Sargis 2008
  Lagomorpha
    61 S Brachylagus idahoensis Leporidae 38628 F Nowak 1999
  62 S Brachylagus idahoensis Leporidae 38631 M Nowak 1999
  63 S Lepus americanus Leporidae 21111 F Nowak 1999
  64 S Lepus americanus Leporidae 33263 M Nowak 1999
  65 Sf Ochotona princeps Ochtonidae 18435 F Nowak 1999
  66 Sf Ochotona princeps Ochtonidae 60068 M Nowak 1999
  Macroscelidea
    67 S Petrodromus tetradactylus Macroscelididae S2008 U Salton and Sargis 2008
  68 T Elephantulus rufescens Macroscelididae 35475 F Nowak 1999
  69 T Elephantulus rufescens Macroscelididae 34189 M Nowak 1999
  Primates
    70 A Callithrix pygmaea Cebidae 39003 F Nowak 1999
  71 A Callithrix pygmaea Cebidae 39005 M Nowak 1999
  72 A Leontopithecus rosalia Cebidae 75541 F Nowak 1999
  73 A Saguinus oedipus Cebidae 35406 F Nyakatura et al. 2008
  74 A Saguinus oedipus Cebidae 35405 M Nyakatura et al. 2008
  75 A Saimiri sciureus Cebidae 82302 F Nowak 1999
  76 A Saimiri sciureus Cebidae 39014 M Nowak 1999
  Rodentia
    77 F Aplodontia rufa Aplodontiidae 34071 F Samuels and Van Valkenburgh 2008
    78 F Aplodontia rufa Aplodontiidae 34058 M Samuels and Van Valkenburgh 2008
    79 Sa Castor canadensis Castoridae 34116 F Nowak 1999
    80 Sa Castor canadensis Castoridae 34588 M Nowak 1999
    81 T Cavia porcellus Caviidae 72830 F Nowak 1999
    82 T Cavia porcellus Caviidae 72831 M Nowak 1999
    83 F Microtus pennsylvanicus Cricetidae 76622 F Nowak 1999
    84 F Microtus pennsylvanicus Cricetidae 34326 M Nowak 1999
    85 Sa Ondatra zibethicus Cricetidae 34324 F Howell 1930; Gingerich 2003
    86 Sa Ondatra zibethicus Cricetidae 72879 M Howell 1930; Gingerich 2003
    87 Sf Synaptomys borealis Cricetidae 66559 F Nowak 1999
    88 Sf Synaptomys borealis Cricetidae 66561 M Nowak 1999
    89 S Zapus princeps Dipodidae 76721 F Nowak 1999
    90 S Zapus princeps Dipodidae 75136 M Nowak 1999
    91 Sc Coendou prehensilis Erethizontidae 34176 F Nowak 1999
    92 F Thomomys bottae Geomyidae 44624 F Nowak 1999
    93 F Thomomys bottae Geomyidae 38261 M Nowak 1999
    94 F Chaetodipus fallax Heteromyidae 47378 F Lackey 1996
    95 F Chaetodipus fallax Heteromyidae 47379 M Lackey 1996
    96 S Dipodomys deserti Heteromyidae 78740 F Samuels and Van Valkenburgh 2008
    97 Sf Octodon degus Octodontidae 48984 F Nowak 1999
    98 Sf Octodon degus Octodontidae 39501 M Nowak 1999
    99 A Sciurus aberti Sciuridae 35278 F Nowak 1999
    100 A Sciurus aberti Sciuridae 82349 M Nowak 1999
    101 A Sciurus carolinensis Sciuridae 20646 F Nowak 1999
    102 A Sciurus carolinensis Sciuridae 30010 M Nowak 1999
    103 G Glaucomys sabrinus Sciuridae 36334 F Samuels and Van Valkenburgh 2008
    104 G Glaucomys sabrinus Sciuridae 35129 M Samuels and Van Valkenburgh 2008
    105 G Glaucomys volans Sciuridae 43897 F Nowak 1999
    106 G Glaucomys volans Sciuridae 32254 M Nowak 1999
    107 Sf Cynomys ludovicianus Sciuridae 75774 M Nowak 1999
  Scandentia
    108 Sc Tupaia glis Tupaiidae 34227 F Van Valkenburgh 1987
    109 Sc Tupaia glis Tupaiidae 34225 M Van Valkenburgh 1987

 

 

TABLE 5. Landmark number, location, type, and source reference for two-dimensional geometric morphometrics (2D GM) points on therian distal humeri. All points were digitized on distal images of specimens; see Figure 3 for more details. Numbers in parentheses are the landmark number for the point within the referenced source. Abbreviations: LM, landmark; Type, landmark type according to Zelditch et al. 2004.

LM Location Type Source Reference
1 Lateral point on the posterior trochlea II Steiner-Souza et al. 2010
2 Medial point on the posterior trochlea II Steiner-Souza et al. 2010
3 Medial point on the anterior trochlea II Steiner-Souza et al. 2010
4 Lateral point on the anterior capitulum II Steiner-Souza et al. 2010
5 Most lateral point on the ectepicondyle III Schutz and Guralnick 2007 (5); Steiner-Souza et al. 2010
6 Most medial point on the entepicondyle III Milne et al. 2009 (12); Steiner-Souza et al. 2010