Project 2292: Z. Luo, S. M. Gatesy, F. A. Jenkins, W. W. Amaral, N. H. Shubin. 2015. Mandibular and dental characteristics of Late Triassic mammaliaform Haramiyavia and their ramifications for basal mammal evolution. Proceedings of the National Academy of Sciences. 112 (51):E7101-E7109.

Specimen: † Haramiyavia clemmenseni Jenkins, F. A. Jr. et al, 1997 (MCZ/7:G95)
View: skull
View: skull
Abstract
Haramiyavia clemmenseni from the Rhaetic (Late Triassic) of East Greenland, as one of the earliest-known mammaliaforms, has held an important place in understanding the timing of the earliest radiation of the group. Reanalysis of the type specimen, using high resolution computed tomography (μCT), has revealed new details, such as the presence of the dentary condyle of the mammalian jaw hinge, and the postdentary trough for mandibular attachment of the middle ear - a transitional condition of the predecessors to crown Mammalia. Our tests of competing phylogenetic hypotheses with these new data show that Late Triassic haramiyids are a separate clade from multituberculate mammals, and are excluded from the Mammalia. Consequently, hypotheses of a Late Triassic diversification of the Mammalia, which are dependent on multituberculate affinities of haramiyidans, are rejected. Scanning Electron Microscopy (SEM) study of tooth wear facets and kinematic functional simulation of occlusion with virtual 3D models from CT scans confirm that Haramiyavia, had a major orthal occlusion with the tallest lingual cusp of the lower occluding into the lingual embrasure of the upper molars, followed by a short palinal movement along the cusp rows alternating between upper and lower molars. This differs from the minimal orthal, but extensive palinal, occlusal movement of multituberculate mammals, previously regarded as relatives of haramiyidans. The disparity of tooth morphology and the diversity of dental functions of haramiyids and their contemporary mammaliaforms suggest dietary diversification is a major factor in the earliest mammaliaform evolution.Read the article »
Article DOI: 10.1073/pnas.1519387112
Project DOI: 10.7934/P2292, http://dx.doi.org/10.7934/P2292
This project contains | Matrices |
---|---|
Download Project SDD File ![]() | Total scored cells: 29851 Total media associated with cells: 0 Total labels associated with cell media: 0 |
Characters | |
Total characters: 497 Total characters with associated media: 0 Total characters with media with labels: 0 Total character states: 1329 Total character states with associated media: 0 Total character states with media with labels:0 Total unordered/ordered characters:497/0 |
Currently Viewing:
MorphoBank Project 2292
MorphoBank Project 2292
- Creation Date:
19 October 2015 - Publication Date:
03 December 2015 - Project views: 51692
- Media downloads: 20
- Matrix downloads: 48
Authors' Institutions
- Brown University
- Harvard University
- University of Chicago
Members
member name | taxa ![]() |
specimens ![]() |
media ![]() | chars ![]() | character
| cell scorings (scored, NPA, "-") | cell
| rules ![]() | ||||||
Zhe-Xi Luo Project Administrator | 1 | 2 | 5 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
Maureen Admin Full membership ![]() | 114 | 0 | 0 | 497 | 0 | 0 | 29851 (29843, 0, 8) | 0 | 0 | 0 | ||||
MorphoBank Curator Full membership ![]() | 0 | 2 | 5 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 |
Taxonomic Overview for Matrix 'M23636' (114 Taxa)
taxon | unscored cells |
scored cells ![]() |
no cell support ![]() |
NPA cells |
"-" cells | cell images | labels on cell images |
member access |
[1] † Thrinaxodon Last Modified in 12/02/15 | 171 | 326 | 326 | 0 | 0 | 0 | 0 | 3 ![]() |
[2] † Massetognathus Last Modified in 12/02/15 | 235 | 262 | 262 | 0 | 0 | 0 | 0 | 3 ![]() |
[3] † Probainognathus Last Modified in 12/02/15 | 256 | 241 | 241 | 0 | 0 | 0 | 0 | 3 ![]() |
[4] † Tritylodontids Last Modified in 12/02/15 | 138 | 362 | 359 | 0 | 0 | 0 | 0 | 3 ![]() |
[5] † Pachygenelus Last Modified in 12/02/15 | 201 | 296 | 296 | 0 | 0 | 0 | 0 | 3 ![]() |
[6] † Adelobasileus Last Modified in 12/02/15 | 405 | 92 | 92 | 0 | 0 | 0 | 0 | 3 ![]() |
[7] † Sinoconodon Last Modified in 12/02/15 | 164 | 334 | 333 | 0 | 0 | 0 | 0 | 3 ![]() |
[8] † Morganucodon Last Modified in 12/02/15 | 111 | 394 | 386 | 0 | 0 | 0 | 0 | 3 ![]() |
[9] † Megazostrodon Last Modified in 12/02/15 | 139 | 358 | 358 | 0 | 0 | 0 | 0 | 3 ![]() |
[10] † Haldanodon Last Modified in 12/02/15 | 184 | 313 | 313 | 0 | 0 | 0 | 0 | 3 ![]() |
[11] † Castorocauda Last Modified in 12/02/15 | 280 | 217 | 217 | 0 | 0 | 0 | 0 | 3 ![]() |
[12] † Megaconus Last Modified in 12/02/15 | 283 | 214 | 214 | 0 | 0 | 0 | 0 | 3 ![]() |
[13] † Eleutherodon Last Modified in 12/02/15 | 448 | 49 | 49 | 0 | 0 | 0 | 0 | 3 ![]() |
[14] † Sineleutherus Last Modified in 12/02/15 | 442 | 56 | 55 | 0 | 0 | 0 | 0 | 3 ![]() |
[15] † Thomasia Last Modified in 12/02/15 | 434 | 63 | 63 | 0 | 0 | 0 | 0 | 3 ![]() |
[16] † Haramiyavia Last Modified in 12/02/15 | 379 | 118 | 118 | 0 | 0 | 0 | 0 | 3 ![]() |
[17] † Arboroharamiya Last Modified in 12/02/15 | 345 | 153 | 152 | 0 | 0 | 0 | 0 | 3 ![]() |
[18] † Xianshou linglong Last Modified in 12/02/15 | 289 | 208 | 208 | 0 | 0 | 0 | 0 | 3 ![]() |
[19] † Xianshou songae Last Modified in 12/02/15 | 295 | 202 | 202 | 0 | 0 | 0 | 0 | 3 ![]() |
[20] † Shenshou Last Modified in 12/02/15 | 236 | 262 | 261 | 0 | 0 | 0 | 0 | 3 ![]() |
[21] † Rugosodon Last Modified in 12/02/15 | 252 | 245 | 245 | 0 | 0 | 0 | 0 | 3 ![]() |
[22] † Kuehneodon Last Modified in 12/02/15 | 369 | 128 | 128 | 0 | 0 | 0 | 0 | 3 ![]() |
[23] † Sinobaatar Last Modified in 12/02/15 | 230 | 268 | 267 | 0 | 0 | 0 | 0 | 3 ![]() |
[24] † Plagiaulacids Last Modified in 12/02/15 | 266 | 236 | 231 | 0 | 0 | 0 | 0 | 3 ![]() |
[25] † Cimolodontans Last Modified in 12/02/15 | 100 | 419 | 397 | 0 | 0 | 0 | 0 | 3 ![]() |
[26] † Hadrocodium Last Modified in 12/02/15 | 242 | 255 | 255 | 0 | 0 | 0 | 0 | 3 ![]() |
[27] † Shuotherium Last Modified in 12/02/15 | 376 | 121 | 121 | 0 | 0 | 0 | 0 | 3 ![]() |
[28] † Pseudotribos Last Modified in 12/02/15 | 309 | 188 | 188 | 0 | 0 | 0 | 0 | 3 ![]() |
[29] † Asfaltomylos Last Modified in 12/02/15 | 400 | 97 | 97 | 0 | 0 | 0 | 0 | 3 ![]() |
[30] † Ambondro Last Modified in 12/02/15 | 421 | 76 | 76 | 0 | 0 | 0 | 0 | 3 ![]() |
[31] † Ausktribosphenos Last Modified in 12/02/15 | 382 | 115 | 115 | 0 | 0 | 0 | 0 | 3 ![]() |
[32] † Bishops Last Modified in 12/02/15 | 381 | 116 | 116 | 0 | 0 | 0 | 0 | 3 ![]() |
[33] † Teinolophos Last Modified in 12/02/15 | 405 | 92 | 92 | 0 | 0 | 0 | 0 | 3 ![]() |
[34] † Steropodon Last Modified in 12/02/15 | 423 | 74 | 74 | 0 | 0 | 0 | 0 | 3 ![]() |
[35] † Obdurodon Last Modified in 12/02/15 | 259 | 238 | 238 | 0 | 0 | 0 | 0 | 3 ![]() |
[36] † Ornithorhynchus Last Modified in 12/02/15 | 130 | 369 | 367 | 0 | 0 | 0 | 0 | 3 ![]() |
[37] † Tachyglossus Last Modified in 12/02/15 | 178 | 321 | 319 | 0 | 0 | 0 | 0 | 3 ![]() |
[38] † Fruitafossor Last Modified in 12/02/15 | 310 | 187 | 187 | 0 | 0 | 0 | 0 | 3 ![]() |
[39] † Gobiconodon Last Modified in 12/02/15 | 127 | 371 | 370 | 0 | 0 | 0 | 0 | 3 ![]() |
[40] † Repenomamus Last Modified in 12/02/15 | 119 | 378 | 378 | 0 | 0 | 0 | 0 | 3 ![]() |
[41] † Amphilestes Last Modified in 12/02/15 | 399 | 98 | 98 | 0 | 0 | 0 | 0 | 3 ![]() |
[42] † Yanoconodon Last Modified in 12/02/15 | 198 | 299 | 299 | 0 | 0 | 0 | 0 | 3 ![]() |
[43] † Jeholodens Last Modified in 12/02/15 | 182 | 315 | 315 | 0 | 0 | 0 | 0 | 3 ![]() |
[44] † Trioracodon Last Modified in 12/02/15 | 343 | 155 | 154 | 0 | 0 | 0 | 0 | 3 ![]() |
[45] † Priacodon Last Modified in 12/02/15 | 297 | 201 | 200 | 0 | 0 | 0 | 0 | 3 ![]() |
[46] † Tinodon Last Modified in 12/02/15 | 396 | 102 | 101 | 0 | 0 | 0 | 0 | 3 ![]() |
[47] † Akidolestes Last Modified in 12/02/15 | 205 | 292 | 292 | 0 | 0 | 0 | 0 | 3 ![]() |
[48] † Spalacotherium Last Modified in 12/02/15 | 344 | 153 | 153 | 0 | 0 | 0 | 0 | 3 ![]() |
[49] † Zhangheotherium Last Modified in 12/02/15 | 142 | 355 | 355 | 0 | 0 | 0 | 0 | 3 ![]() |
[50] † Maotherium Last Modified in 12/02/15 | 140 | 359 | 357 | 0 | 0 | 0 | 0 | 3 ![]() |
[51] † Dryolestes Last Modified in 12/02/15 | 322 | 175 | 175 | 0 | 0 | 0 | 0 | 3 ![]() |
[52] † Henkelotherium Last Modified in 12/02/15 | 252 | 245 | 245 | 0 | 0 | 0 | 0 | 3 ![]() |
[53] † Amphitherium Last Modified in 12/02/15 | 357 | 140 | 140 | 0 | 0 | 0 | 0 | 3 ![]() |
[54] † Peramus Last Modified in 12/02/15 | 333 | 166 | 164 | 0 | 0 | 0 | 0 | 3 ![]() |
[55] † Vincelestes Last Modified in 12/02/15 | 94 | 405 | 403 | 0 | 0 | 0 | 0 | 3 ![]() |
[56] † Nanolestes Last Modified in 12/02/15 | 351 | 146 | 146 | 0 | 0 | 0 | 0 | 3 ![]() |
[57] † Kielantherium Last Modified in 12/02/15 | 356 | 139 | 138 | 0 | 3 | 0 | 0 | 3 ![]() |
[58] † Aegialodon Last Modified in 12/02/15 | 437 | 60 | 60 | 0 | 0 | 0 | 0 | 3 ![]() |
[59] † Montanalestes Last Modified in 12/02/15 | 381 | 116 | 116 | 0 | 0 | 0 | 0 | 3 ![]() |
[60] † Prokennalestes Last Modified in 12/02/15 | 279 | 220 | 218 | 0 | 0 | 0 | 0 | 3 ![]() |
[61] † Murtoilestes Last Modified in 12/02/15 | 406 | 91 | 91 | 0 | 0 | 0 | 0 | 3 ![]() |
[62] † Eomaia Last Modified in 12/02/15 | 171 | 326 | 326 | 0 | 0 | 0 | 0 | 3 ![]() |
[63] † Juramaia Last Modified in 12/02/15 | 304 | 188 | 188 | 0 | 5 | 0 | 0 | 3 ![]() |
[64] † Kennalestes Last Modified in 12/02/15 | 194 | 305 | 303 | 0 | 0 | 0 | 0 | 3 ![]() |
[65] † Asioryctes Last Modified in 12/02/15 | 117 | 380 | 380 | 0 | 0 | 0 | 0 | 3 ![]() |
[66] † Ukhaatherium Last Modified in 12/02/15 | 200 | 297 | 297 | 0 | 0 | 0 | 0 | 3 ![]() |
[67] † Zalambdalestes Last Modified in 12/02/15 | 75 | 430 | 422 | 0 | 0 | 0 | 0 | 3 ![]() |
[68] † Daulestes Last Modified in 12/02/15 | 258 | 240 | 239 | 0 | 0 | 0 | 0 | 3 ![]() |
[69] † Aspanlestes Last Modified in 12/02/15 | 358 | 139 | 139 | 0 | 0 | 0 | 0 | 3 ![]() |
[70] † Eoungulatum Last Modified in 12/02/15 | 372 | 125 | 125 | 0 | 0 | 0 | 0 | 3 ![]() |
[71] † Cimolestes Last Modified in 12/02/15 | 328 | 169 | 169 | 0 | 0 | 0 | 0 | 3 ![]() |
[72] † Gypsonictops Last Modified in 12/02/15 | 331 | 166 | 166 | 0 | 0 | 0 | 0 | 3 ![]() |
[73] † Protungulatum Last Modified in 12/02/15 | 276 | 222 | 221 | 0 | 0 | 0 | 0 | 3 ![]() |
[74] Erinaceus Last Modified in 10/22/15 | 40 | 458 | 457 | 0 | 0 | 0 | 0 | 3 ![]() |
[75] † Leptictis Last Modified in 12/02/15 | 76 | 421 | 421 | 0 | 0 | 0 | 0 | 3 ![]() |
[76] Canis Last Modified in 10/22/15 | 51 | 447 | 446 | 0 | 0 | 0 | 0 | 3 ![]() |
[77] Felis Last Modified in 10/22/15 | 56 | 441 | 441 | 0 | 0 | 0 | 0 | 3 ![]() |
[78] Rattus Last Modified in 10/22/15 | 98 | 399 | 399 | 0 | 0 | 0 | 0 | 3 ![]() |
[79] Oryctolagus Last Modified in 10/22/15 | 88 | 409 | 409 | 0 | 0 | 0 | 0 | 3 ![]() |
[80] Bradypus Last Modified in 10/22/15 | 171 | 327 | 326 | 0 | 0 | 0 | 0 | 3 ![]() |
[81] Tamandua Last Modified in 10/22/15 | 204 | 293 | 293 | 0 | 0 | 0 | 0 | 3 ![]() |
[82] † Glyptotherium Last Modified in 12/02/15 | 179 | 318 | 318 | 0 | 0 | 0 | 0 | 3 ![]() |
[83] Dasypus Last Modified in 10/22/15 | 166 | 332 | 331 | 0 | 0 | 0 | 0 | 3 ![]() |
[84] Chaetophractus Last Modified in 10/22/15 | 166 | 333 | 331 | 0 | 0 | 0 | 0 | 3 ![]() |
[85] Euphractus Last Modified in 10/22/15 | 167 | 332 | 330 | 0 | 0 | 0 | 0 | 3 ![]() |
[86] † Holoclemensia Last Modified in 12/02/15 | 399 | 98 | 98 | 0 | 0 | 0 | 0 | 3 ![]() |
[87] † Sinodelphys Last Modified in 12/02/15 | 227 | 270 | 270 | 0 | 0 | 0 | 0 | 3 ![]() |
[88] † Deltatheridium Last Modified in 12/02/15 | 213 | 286 | 284 | 0 | 0 | 0 | 0 | 3 ![]() |
[89] † Atokatheridium Last Modified in 12/02/15 | 407 | 90 | 90 | 0 | 0 | 0 | 0 | 3 ![]() |
[90] † Sulestes Last Modified in 12/02/15 | 312 | 185 | 185 | 0 | 0 | 0 | 0 | 3 ![]() |
[91] † Asiatherium Last Modified in 12/02/15 | 177 | 321 | 320 | 0 | 0 | 0 | 0 | 3 ![]() |
[92] † Kokopellia Last Modified in 12/02/15 | 346 | 152 | 151 | 0 | 0 | 0 | 0 | 3 ![]() |
[93] † Anchistodelphys Last Modified in 12/02/15 | 403 | 94 | 94 | 0 | 0 | 0 | 0 | 3 ![]() |
[94] † Albertatherium Last Modified in 12/02/15 | 437 | 60 | 60 | 0 | 0 | 0 | 0 | 3 ![]() |
[95] † Didelphodon Last Modified in 12/02/15 | 264 | 233 | 233 | 0 | 0 | 0 | 0 | 3 ![]() |
[96] † Pediomys Last Modified in 12/02/15 | 297 | 200 | 200 | 0 | 0 | 0 | 0 | 3 ![]() |
[97] † Turgidodon Last Modified in 12/02/15 | 316 | 181 | 181 | 0 | 0 | 0 | 0 | 3 ![]() |
[98] † Mayulestes Last Modified in 12/02/15 | 101 | 396 | 396 | 0 | 0 | 0 | 0 | 3 ![]() |
[99] † Pucadelphys Last Modified in 12/02/15 | 79 | 418 | 418 | 0 | 0 | 0 | 0 | 3 ![]() |
[100] † Andinodelphys Last Modified in 12/02/15 | 196 | 301 | 301 | 0 | 0 | 0 | 0 | 3 ![]() |
[101] Didelphis Last Modified in 10/22/15 | 37 | 462 | 460 | 0 | 0 | 0 | 0 | 3 ![]() |
[102] Marmosa Last Modified in 10/22/15 | 37 | 460 | 460 | 0 | 0 | 0 | 0 | 3 ![]() |
[103] Caenolestes Last Modified in 10/22/15 | 70 | 427 | 427 | 0 | 0 | 0 | 0 | 3 ![]() |
[104] Dasyurus Last Modified in 10/22/15 | 42 | 456 | 455 | 0 | 0 | 0 | 0 | 3 ![]() |
[105] Perameles Last Modified in 10/22/15 | 65 | 432 | 432 | 0 | 0 | 0 | 0 | 3 ![]() |
[106] Dromiciops Last Modified in 10/22/15 | 45 | 452 | 452 | 0 | 0 | 0 | 0 | 3 ![]() |
[107] Thylacomyidae Last Modified in 10/22/15 | 146 | 351 | 351 | 0 | 0 | 0 | 0 | 3 ![]() |
[108] Macropus Last Modified in 10/22/15 | 72 | 427 | 425 | 0 | 0 | 0 | 0 | 3 ![]() |
[109] Acrobates Last Modified in 10/22/15 | 74 | 423 | 423 | 0 | 0 | 0 | 0 | 3 ![]() |
[110] Phascolarctos Last Modified in 10/22/15 | 75 | 422 | 422 | 0 | 0 | 0 | 0 | 3 ![]() |
[111] Vombatus Last Modified in 10/22/15 | 75 | 422 | 422 | 0 | 0 | 0 | 0 | 3 ![]() |
[112] Phalanger Last Modified in 10/22/15 | 69 | 428 | 428 | 0 | 0 | 0 | 0 | 3 ![]() |
[113] Pseudocheirus Last Modified in 10/22/15 | 70 | 427 | 427 | 0 | 0 | 0 | 0 | 3 ![]() |
[114] Petauroides Last Modified in 10/22/15 | 71 | 426 | 426 | 0 | 0 | 0 | 0 | 3 ![]() |
Project views 
type | number of views | Individual items viewed (where applicable) |
Total project views | 51692 | |
Project overview | 5076 | |
Media views | 12139 | Media search (4747 views); M406226 (768 views); M406227 (779 views); M406228 (734 views); M406245 (789 views); M402214 (748 views); M402215 (673 views); M406231 (669 views); M406246 (785 views); M406232 (699 views); M406248 (748 views); |
Documents list | 1615 | |
Matrix views | 2372 | Matrix landing page (1939 views); Matrix (433 views); |
Taxon list | 21501 | |
Specimen list | 4182 | |
Views for media list | 3712 | |
Bibliography | 1095 |
Project downloads 
type | number of downloads | Individual items downloaded (where applicable) |
Total downloads from project | 494 | |
Media downloads | 20 | M406228 (2 downloads); M402215 (1 download); M406226 (6 downloads); M406245 (6 downloads); M406246 (3 downloads); M406248 (2 downloads); |
Project downloads | 419 | |
Matrix downloads | 48 | Matrix (48 downloads); |
Document downloads | 7 | Luo ZX et al. (2015) PNAS List of Characters accompanying SI Appendix (5, 6, & 7) (7 downloads); |