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,QApatosaurus/$*VZHUHQRWGHSRVLWHGXQWLODQDQLPDOUHDFKHGDGXOWVL]HEHIRUHZKLFKD]RQDO)/%GRPLQDWHGLQWKHFRUWH[ &XUU\ 7KHKDGURVDXULGMaiasauraOLNHO\JUHZWRPLQLWVILUVW\HDUZLWKRQVHWRIVRPDWLFPDWXULW\RFFXUULQJE\DJH +RUQHUHWDO 7KHDJHZKLFKWKLVGHFUHDVHLQJURZWKUDWHRFFXUVLQPsittacosaurusKDVEHHQHVWLPDWHGD\HDUV (ULFNVRQDQG7XPDQRYD (VWLPDWHVIRUVDXULVFKLDQWD[DLQFOXGHWKHVPDOOWKHURSRGVCoelophysis ±\HDUV&KLQVDP\ DQGTroodon ±\HDUV9DUULFFKLR
(ULFNVRQHWDO WKHVDXURSRGVJanenschia \HDUV6DQGHU DQGLapparentosaurus\HDUVGH5LFTOpV Tyrannosaurus \HDUV(ULFNVRQHWDO DQGWKH$OEHUWDQW\UDQQRVDXULGVAlbertosaurusDaspletosaurusDQGGorgosaurus ±\HDUV(ULFNVRQHWDO66
 5HODWLYHO\PRGHUDWHJURZWKWUDMHFWRULHVDUHJHQHUDOO\DVVRFLDWHGZLWKUHODWLYHO\VPDOORUQLWKLVFKLDQVVXFKDVWKH³K\SVLORSKRGRQWLG´Orodromeus +RUQHUHWDO ,QMXYHQLOHPinacosaurusVSHFLPHQVWKHKLVWRORJLFDOIHDWXUHVRIWKHOLPEERQHVLQGLFDWHUDSLGO\JURZLQJERQHOLNHWKDWUHSRUWHGIRURWKHUMXYHQLOHGLQRVDXUV 6DQGHU+RUQHUHWDO+D\DVKLHWDO 1RQHRIWKHGastoniaHOHPHQWVVHFWLRQHGQRWHYHQWKHODUJHVWIRUHDFKHOHPHQWW\SHH[KLELWZKDWPD\XQHTXLYRFDOO\UHIHUUHGWRDVDQH[WHUQDOIXQGDPHQWDOV\VWHP ()6 7KHUHIRUHQRQHRIWKHVSHFLPHQVPD\EHVDLGWRKDYHUHDFKHGVRPDWLFPDWXULW\E\WKHWLPHRIGHDWK1HYHUWKHOHVVLWLVFOHDUWKDWVHFRQGDU\UHPRGHOLQJLVH[WHQVLYHLQWKHODUJHU
HOHPHQWV7KLVPD\PHDQWKDWDQN\ORVDXUVH[KLELWHGDPRUHSURORQJHGSHULRGRIJURZWKUDWH
FKDQJHIURPDPD[LPDOUDWHLQMXYHQLOHVWRDFHVVDWLRQRIJURZWKDWPD[LPDOVL]H7KLVLV
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SURPLQHQWVLJPRLGDOJURZWKFXUYHK\SRWKHVL]HGIRURWKHUGLQRVDXUVVXFKDVW\UDQQRVDXULGV(ULFNVRQHWDO 1HYHUWKHOHVVDQN\ORVDXUVOLNHStegosaurusVWLOOKDGKLJKHUPD[LPDOMXYHQLOHJURZWKUDWHVWKDQWKHEDVDOWK\UHRSKRUDQVScutellosaurusLQZKLFKWKHOLPEERQHVDUH
FKDUDFWHUL]HGE\VORZIRUPLQJ]RQDOSDUDOOHOILEUHHGWRODPHOODUERQH 3DGLDQHWDO 7LPLQJRI2VWHRGHUP'HYHORSPHQW
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RVVLFOHVFDQQRWEHXVHGDVDQLQGHSHQGHQWVRXUFHRIH[DFWDJHGDWDLQDQN\ORVDXUVDWSUHVHQW,Q
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67PDWHULDOLQPinacosaurusDVLPLODUGHOD\KDVEHHQK\SRWKHVL]HGIRUDQN\ORVDXUVDVZHOO %XUQV
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+DYHUVLDQERQHRIODUJH !PD[LPXPVL]H DQN\ORVDXUOLPEERQHV7KHILEUHVRFFXULQ
EXQGOHVDQGWHQGWREOXUWKHRWKHUZLVHFULVSK\SHUPLQHUDOL]HGUHVRUSWLRQOLQHVGHPDUFDWLQJWKHSHULSKHU\RIVHFRQGDU\RVWHRQV,QDVPDOOHUUDGLXV PD[LPXPNQRZQVL]H WKHVHILEUHVDUH
68SUHVHQWEXWDUHPRUHVSDUVHWKDQLQWKHODUJHUHOHPHQWV 6WHLQHWDO 6XFKVWUXFWXUHVZHUHQRWREVHUYHGKHUHLQHLWKHUGastoniaRUPinacosaurus,WLVSRVVLEOHWKDWWKHLQFRUSRUDWLRQRIWKHVHILEUHVLQWRWKHUHPRGHOHGERQHPDWUL[VHUYHGWRHQKDQFHWKHVWUHQJWKRIWKH+DYHUVLDQ
ERQHDVK\SRWKHVL]HGE\6WHLQHWDO  7KHDEVHQFHRIWKLVIHDWXUHLQGastoniaKRZHYHUZKLFKLVNQRZQWRKDYHKDGIXOO\GHYHORSHGRVWHRGHUPVE\WKHVXEDGXOWVWDJHZRXOGQRWOHQGVXSSRUWWRWKDWK\SRWKHVLV1HYHUWKHOHVVLWLVSRVVLEOHWKDWWKHVHILEUHVDUHFKDUDFWHULVWLFRIWKH
ODWHVWDGXOWRQWRJHQHWLFVWDJHVLQDQN\ORVDXUVDVVRFLDWHGZLWKVHYHUDOJHQHUDWLRQVRIVHFRQGDU\RVWHRQV

697$%/((OHPHQWVIURPMXYHQLOHVSHFLPHQVRIPinacosaurus grangeriIURP$ODJ7HHJKLVWRORJLFDOO\VDPSOHG/HQJWKLVLQPP
Inst.
HMNS
HMNS
HMNS
HMNS
HMNS
HMNS
HMNS
HMNS
MPCͲD
MPCͲD
MPCͲD
MPCͲD
MPCͲD

Number95Ͳ11Ͳ5
95Ͳ11Ͳ5
95Ͳ11Ͳ5
95Ͳ11Ͳ25
95Ͳ11Ͳ65
95Ͳ11Ͳ65
00Ͳ11Ͳ1
00Ͳ11Ͳ1
100/1322
100/1322
100/1322
100/1344
100/1344

Element
Humerus
Femur
RadiusFibulaHumerusRadiusFemur
Radius
Femur
Femur
Femur
Femur
Femur

Length
150
180
100
131
85
220
97
198
210
216
169
186

Side

R
R
R
L
L
L
R





70
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UHIHUVWRWKHPD[LPXPQXPEHURI/$*VLGHQWLILDEOHLQHDFKVHFWLRQ/HQJWKLVLQPP
DMNH Element Length Side LAGs
50082 Femur
509
R
7
50215 Femur
414
L
2
53703 Femur
437
L
5
53705 Femur
390
L
350234 Fibula
210
R
1
50238 Fibula
227
R
052077 Fibula
248
L
2
53858 Fibula
195
L
1
56725 Fibula
239
R
053338 Humerus 297
R
453339 Humerus 326
R557651 Humerus 364
R61109 Humerus 276
L350302 Radius
304
450305 Radius
151
550328 Radius
154
R
4
53084 Radius
167
253283 Radius
140
150114 Tibia
243
L
650275 Tibia
294
L
4
51861 Tibia
252
R
1
57685 Tibia
461133 Tibia
206
R
250265 Ulna
150
L
350326 Ulna
195
R
2
51965 Ulna
184
L
3
53089 Ulna
173
L
361129 Ulna
218
L
650110 Humerus 234
R
3
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edwardsorumLQZKLFKWKHJUHDWHUWURFKDQWHULVORFDWHGPRUHSUR[LPDOO\ &DUSHQWHU 7KH
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KDOIULQJLQWKHVDPHWD[RQLQWKDWWKH\DUHSRO\JRQDOKRZHYHUWKHPHGLDQNHHOVGLYHUJHPRUH
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364 280 84 302 138 100 119 292
477 317 160 353 171 81 138 328
455 293 162 332 167  133 312'01+

ROM1215
TMP00.12.158
TMP81.3.3
TMP83.25.2
TMP84.0.32
TMP97.9.1
USNM11868

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CMN8879   

169

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     

35&01

203 86&01

Pawpawsaurus 245 164 80

6SHFLPHQ

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Pawpawsaurus 0.46 0.45 0.47 0.52 0.45 0.25 0.32 0.51&01
0.65 0.63 0.74 0.72 0.62 0.57 0.63 0.84&01
0.83 0.73 0.76 0.79 0.94 0.39 0.67 0.84
CMN8879
0.80 0.85 0.69 0.80 0.87 0.42 1.00 0.96
'01+
0.91 0.97 0.78 0.56 1.00 0.61 0.72 0.52
ROM1215
0.76 0.75 0.80 0.77 0.97 0.40 0.59 0.83
TMP00.12.158 1.00 1.00 1.00 0.86 0.67 0.61 0.86 0.98
TMP81.3.3
0.81 0.73 0.98 1.00 0.75 0.55 0.65 0.93
TMP83.25.2
0.83 0.77 0.95 0.71 0.79 1.00 0.81 0.85
TMP84.0.32
0.68 0.77 0.49 0.77 0.72 0.73 0.67 0.89
TMP97.9.1
0.89 0.87 0.93 0.90 0.90 0.59 0.78 1.00
USNM11868
0.85 0.81 0.94 0.85 0.88 0.50 0.75 0.95

6SHFLPHQ

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7$%/(&UDQLDOTXDQWLWDWLYHGDWDIRU/DWH&UHWDFHRXVQRGRVDXULGVSHFLPHQVDQGWKHKRORW\SH 608 RIWKHRXWJURXS

Scelidosaurus
Emausaurus
Pawpawsaurus
AMNH3076
AMNH5381
AMNH5665
BHI6225
BHI6332
CMN2759
CMN8531
DMNH468
DPMWA90Ͳ25
ROM1215
ROM20892
TMP00.12.158
TMP83.25.2
TMP98.67.1
TMP98.98.1
UALVP16249
USNM11868

VRPHQHZFKDUDFWHUVDUHEDVHGRQ%DNNHU  DQG&DUSHQWHU  

Emausaurus, PawpawsaurusDQGScelidosaurus6RPHFUDQLDODQGGHQWDOFKDUDFWHUVDUHPRGLILHGIURP9LFNDU\RXVHWDO  DQG

7$%/(&KDUDFWHUVSHFLPHQPDWUL[XVHGLQSDUVLPRQ\DQDO\VLVRI/DWH&UHWDFHRXV1RGRVDXULGDHDQGWKUHHRXWJURXSWD[D

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137

0 ? 0 ? ? 0 0 ? 0 0 ? ? 0 0 0 0 ? 0 0 0 0 0 0 0 0 1 0 0 0 0 ? ? ? ? ? 0 0 0 ? ? ? 0
0 0 0 0 ? 0 ? 0 ? ? ? ? ? ? ? 0 ? 0 0 0 0 0 0 0 0 1 0 0 0 0 ? ? ? 0 ? 0 0 0 ? ? ? 0
0 0 0 1 1 1 1 1 1 1 1 2 1 1 1 1 ? 1 1 0 1 1 1 1 1 ? ? 1 ? 2 0 0 0 ? ? ? 0 1 0 1 1 0? ? ? 1 ? 1 1 ? ? ? ? ? ? ? ? ? ? 1 1 1 1 1 1 ? ? ? ? ? ? ? ? 0 0 ? ? ? 0 4 ? 1 1 ?
1 ? 1 1 0 ? 1 ? 1 ? ? ? 1 ? 1 1 ? 1 1 1 0 1 1 ? 1 ? 1 ? ? 1 ? 1 0 ? ? ? 0 4 ? 0 ? ?
1 ? 1 ? ? 1 1 ? 1 1 1 ? 1 ? 1 1 1 1 1 1 ? 1 ? 1 1 ? 1 1 0 2 ? 1 0 ? 1 1 0 4 0 ? ? ?
? ? ? ? ? ? ? 1 1 1 ? ? ? ? 1 ? ? 1 1 1 1 ? 1 ? ? ? ? ? ? ? ? 1 0 ? 1 ? 0 2 ? ? ? 1? ? 1 1 0 ? ? ? 1 ? 1 ? ? 1 1 0 ? 1 1 1 1 ? ? 1 ? ? ? ? ? 2 ? ? ? 0 ? ? 0 ? ? 0 ? ?
1 1 1 1 0 1 1 1 1 1 1 2 ? 1 1 1 ? 1 1 1 1 1 1 1 1 ? 1 1 1 2 ? 0 1 ? 0 0 0 3 0 1 0 0
2 ? 1 1 0 1 1 ? 1 1 1 ? 1 ? 1 1 ? 1 1 1 1 1 1 1 1 ? 1 0 0 2 2 1 0 ? 1 2 0 3 ? 1 1 0? ? 1 1 ? 1 1 ? 1 1 1 ? ? ? 1 1 ? 1 1 1 1 1 1 1 ? ? ? ? 0 2 ? ? 0 ? ? ? 0 2 ? 1 0 1? ? ? 1 ? ? 1 ? 1 ? ? ? 1 ? ? ? ? ? 1 ? ? ? 1 1 ? ? ? ? ? ? ? ? 1 ? ? ? 1 2 0 ? ? ?
1 0 1 1 0 1 1 ? 1 1 1 2 1 ? 1 1 ? 1 1 1 1 1 1 1 1 ? 1 0 1 1 1 0 1 1 0 0 1 2 0 0 0 0? ? ? ? ? ? ? ? ? ? ? ? 1 ? 1 0 ? 1 1 1 1 1 ? ? ? ? ? ? ? 2 2 0 ? ? ? ? ? 2 ? ? ? ?
1 ? 1 1 0 ? 1 ? 1 ? 1 2 1 ? 1 1 ? 1 1 1 0 1 1 ? ? ? ? ? 1 1 2 1 0 ? ? ? 0 ? 1 1 0 0? ? 1 1 0 ? 1 ? 1 ? 1 2 1 ? 1 1 ? 1 1 1 1 1 1 ? ? ? ? ? 1 2 ? 0 1 ? ? ? 0 ? 1 0 0 1? ? ? ? ? ? 1 ? 1 ? ? ? ? ? ? ? ? 1 1 1 ? ? 1 ? 1 ? 1 ? ? ? ? 0 0 ? 0 ? 0 ? ? 0 ? 0
0 0 1 1 0 ? 1 ? 1 ? 1 2 1 ? ? ? ? 1 1 1 1 1 1 1 1 ? 1 1 ? ? ? 0 0 ? ? ? 0 4 ? 1 ? 1
0 0 1 1 0 ? 1 1 1 1 ? 2 1 ? 1 1 ? 1 1 1 1 1 1 ? ? ? ? ? ? 2 ? 1 0 ? ? ? 0 ? 1 0 1 1? ? 1 1 0 ? 1 ? 1 ? 1 2 1 ? 1 1 ? 1 1 1 ? 1 1 ? 1 ? 1 1 ? 1 ? 1 0 ? 1 1 0 ? 0 0 0 0

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42

7$%/(/LPEERQHPHDVXUHPHQWV PP IRUVRPH/DWH&UHWDFHRXVQRGRVDXULGVSHFLPHQV$EEUHYLDWLRQV&PLGVKDIWFLUFXPIHUHQFH':PD[LPXPGLVWDOZLGWK/PD[LPXPSUR[LPRGLVWDOOHQJWK0PDQXDO0&PHWDFDUSDO03KPDQXDOSKDODQ[07PHWDWDUVDO
0:PD[LPXPPLGVKDIWZLGWK3SHGDO3:PD[LPXPSUR[LPDOZLGWK6VLGH6SHF
VSHFLPHQQXPEHU
Spec.
S L
PW MW DW C
AMHH5381 humerus
L 563 329 89 219 288
CMN2759 fibula
L 315 17 40 70 106
humerusL 440 244 71 195 201
MungualIͲ2 L 50 38 36 22 99
MungualIIͲ3 L 52 47 42 31 101
MungualIIIͲ3 L
42 31 101
MCI
L 97 53 31 45
MCII
L 110 48 35 51 97
MCIII
L 117 69 37 43 104
MPhIͲ1
L 51 42 37 41
MPhIIͲ1
L 26 55 52 49 129
MPhIIͲ2
L 27 47 49 48 117
MPhIIIͲ1
L 28 53
MPhIIIͲ2
L 22 45 48 47
MT
134 43 35 67 118
Pungual
82 58 50 28 117Pungual
81 54 52 29 120
tibia
L 381 47 70 147 191
tibiaR
39 64 157 193
CMN8531 femur
R 661 244 109 221 283
humerusL 485 195 106 225 266
radius
L 291 112 34 62 164
radius
R 287 53 33 65 181
tibia
R
106
ulna
L 323 194 74 53 207
TMP98.98.1 fibula
R 133 128 50 66 133
humerusR 586 290 89 194 305
radiusR 425 233 109 81 251
ulnaR
69 157 193 

138
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7$%/( QH[WSDJH 9HUWHEUDOPHDVXUHPHQWV PP IRUVRPH/DWH&UHWDFHRXVQRGRVDXULGVSHFLPHQV$EEUHYLDWLRQV$+DQWHULRUFHQWUXPKHLJKW$:DQWHULRUFHQWUXPZLGWK&G
FDXGDO&YFHUYLFDO'GRUVDO+WRWDOKHLJKW/WRWDOFHQWUXPDQWHURSRVWHULRUOHQJWK0:
PLGGOHFHQWUXPZLGWK1&+QHXUDOFDQDOKHLJKW1&:QHXUDOFDQDOZLGWK16+QHXUDOVSLQHKHLJKWPHDVXUHGIURPWRSRIFHQWUXP3+SRVWHULRUFHQWUXPKHLJKW3:SRVWHULRUFHQWUXPZLGWK6VLGH6±VDFUDOV6SHFVSHFLPHQQXPEHU

139
Spec.
SL
H AW MW PW AH PH NCH NCW NSH
AMNH5381 Cd 59 217 93 83 107 80 90 44 21
127D
263 306 136 93 121 115 110 35 23
201
D
93 304 116 61 130 132 115 51 19
189D
81 302 109 75 109 107 104 31 21
181
D89
117 86 130 100 114 40 32
D74
104 153 53 109 119 37
CMN2759 Cd 58
92 83 95
40 25
Cd 51
84 64 83 65 68
23
Cv1 89 77 72
47
28 48
40
Cv2 53 114
51
Cv3 63 113
87
49
76
DS1
87
87
CMN8531 Cd 44 202 80 78 93 82 112 35 18
96
Cd 41
87 81 93 70 76 19 2277
Cd 55 195 102 94 98 75 91 39 25
104
Cd 48
89 84 92 74 80 22 1574
Cd 63 179 107 96 103 75 80 17 1796
Cd 43 202 99 86 102 85 103 32 21
100
Cd1
99
Cv 53 187 98 89 111 79 89 31 48
106D75
106 87 107 69 104 23 39
D
87 240 88 55 85 98 99 30 19
168
D
84 252 82 61 98 91 94 28 28
172
D
73 257 83 55 91 97 91 25 21
166
D
74 247 109 76 117 102 99 29 29
163
D
80 260 103 78 118 98 94 35 26
161
D73
122 96 118
D82
115 91 112
D92
100 88 102
S1
126
154
S2
154
151
S3
151
111
S4 74
112 92 97
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140
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141
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142
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143
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144

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145

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146

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148
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149
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150
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158
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159
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o Gargoyleosaurus &DUSHQWHU0LOHVDQG&ORZDUG
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270
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272
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273
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O 67587+,26$85,1$(123&6$
 Anoplosaurus 6HHOH\
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:LHUVPD
x Europelta carbonensis.LUNODQG$OFDOD/RHZHQ(VStOH]
0DPSHODQG:LHUVPDW\SHVSHFLHV275
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x = Danubiosaurus %XQ]HO
x = Hoplosaurus 6HHOH\
x = Leipsanosaurus 1RSVFD
x = Pleuropeltis 6HHOH\ SDUWLPx = Rhodanosaurus 1RSVFD
x Struthiosaurus austriacus %XQ]HOW\SHVSHFLHVo 3,8:/DWH&UHWDFHRXV ORZHU&DPSDQLDQ *RVDX)RUPDWLRQ *UQEDFK%DVLQ *XWH+RIIQXQJ

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o 1+08./DWH&UHWDFHRXV ODWH
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o 802/9'$±*&9/DWH&UHWDFHRXV&DPSDQLDQ 8QQDPHGXQLW+pUDXOW)UDQFH

277
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2[IRUG&OD\ SUREDEO\Kosmoceras jason =RQH EULFNSLWDW)OHWWRQ&DPEULGJHVKLUH(QJODQG
o Texasetes &RRPEV
 Texasetes pleurohalio &RRPEVW\SHVSHFLHV
x 8610(DUO\&UHWDFHRXV ODWH$OELDQ 3DZ3DZ
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Acanthopholiseucerus6HHOH\
Acanthopholishorridus+X[OH\
Acanthopholismacrocercus6HHOH\
Acanthopholisplatypus6HHOH\ SDUWLP 
Acanthopholisstereocercus6HHOH\ SDUWLP 
Anoplosaurusmajor6HHOH\ SDUWLP 
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Cryptodracoeumerus 6HHOH\ 
Cryptosauruseumerus6HHOH\
Dysganusbicraniatus&RSH
Dysganusencaustus&RSH
Dysganusnaydenianus&RSH
Dysganuspeiganus&RSH
Eucercosaurustanyspondylus6HHOH\
Hierosaurussternbergii:LHODQG
Iguanodonphillipsii6HHOH\
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Onychosaurushungaricus1RSVFD
Paleoscincusafricanus%URRP
Paleoscincuscostatus/HLG\
Paleoscincuslatus0DUVK
Paleoscincusmagoder+HQQLQJ QRPHQQXGXP 
Paranthodonoweni1RSVFD
Peishansaurusphilemys%RKOLQ
Priconodoncrassus0DUVK
Priodontognathusphillipsii 6HHOH\ 
Regnosaurusnorthamptoni0DQWHOO
Rhadinosaurusalcinus6HHOH\
Sauroplitesscutiger%RKOLQ

280
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