Project 5470: M.D. Nelson, O. Lambert, M. D. Uhen. 2025. Reassessment of the iconic Oligo-Miocene heterodont dolphin Squalodon: a redescription of the type species S. grateloupii. Papers in Palaeontology. (In Press)
Abstract
Since its description in 1840, the toothed whale Squalodon was conceptualized as a transitional form between archaeocetes like Basilosaurus and the modern toothed whales (Odontoceti). The cranial morphology of Squalodon approximates that of modern odontocetes; however, their dental patterning more closely resembles that of archaeocetes. Despite representing a transitional form, the Squalodontidae have been neglected in scientific study. The poor taxonomy of the family is the primary cause—significant specimens have not been redescribed, including the type species, S. grateloupii. Complicating matters, the holotype of that species has been missing for over a century, and this calls for the designation of a neotype by the ICZN. We present a neotype for S. grateloupii and synapomorphies for the genus and family which include several features in the posterior basicranium and the periotic; we identified a strongly conserved dental pattern for the Squalodontidae. As the Squalodontidae has been underrepresented in phylogenetic analyses, our analysis included representatives from nearly all clades within Odontoceti, as well as every squalodontid species. Our results demonstrate that the Squalodontidae is monophyletic; unlike previous iterations of this phylogenetic analysis, the Squalodontidae were not recovered in crown Odontoceti. This result supports the hypothesis that homodonty is a synapomorphy of crown Odontoceti and did not evolve convergently, which is supported by four dental synapomorphies for crown odontocetes. Our modern revision of the iconic Squalodon lays the foundation for research into interspecific variation, ontogeny, and biogeography of a key transitional form, so that we better understand the course of odontocete evolution.Project DOI: 10.7934/P5470, http://dx.doi.org/10.7934/P5470
This project contains | Matrices |
---|---|
Download Project SDD File ![]() | Total scored cells: 20396 Total media associated with cells: 0 Total labels associated with cell media: 0 |
Characters | |
Total characters: 295 Total characters with associated media: 0 Total characters with media with labels: 0 Total character states: 769 Total character states with associated media: 0 Total character states with media with labels:0 Total unordered/ordered characters:295/0 |
Currently Viewing:
MorphoBank Project 5470
MorphoBank Project 5470
- Creation Date:
21 August 2024 - Publication Date:
27 January 2025 - Matrix downloads: 1
Authors' Institutions
- Calvert Marine Museum
Members
member name | taxa ![]() |
specimens ![]() |
media ![]() | chars ![]() | character
| cell scorings (scored, NPA, "-") | cell
| rules ![]() | ||||||
Margot Nelson Project Administrator | 104 | 1 | 1 | 295 | 0 | 0 | 20396 (19745, 0, 651) | 0 | 0 | 0 |
Taxonomic Overview for Matrix 'M30097' (104 Taxa)
taxon | unscored cells |
scored cells ![]() |
no cell support ![]() |
NPA cells |
"-" cells | cell images | labels on cell images |
member access |
[1] † Georgiacetus vogtlensis Taxon name last Modified on 01/27/25 | 137 | 146 | 146 | 0 | 12 | 0 | 0 | 1 ![]() |
[2] † Zygorhiza kochii Taxon name last Modified on 01/27/25 | 67 | 220 | 217 | 0 | 11 | 0 | 0 | 1 ![]() |
[3] † Archaeodelphis patrius Taxon name last Modified on 01/27/25 | 174 | 111 | 111 | 0 | 10 | 0 | 0 | 1 ![]() |
[4] † Xenorophus sloanii Taxon name last Modified on 01/27/25 | 196 | 94 | 94 | 0 | 5 | 0 | 0 | 1 ![]() |
[5] † Agorophius pygmaeus Taxon name last Modified on 01/27/25 | 155 | 137 | 136 | 0 | 4 | 0 | 0 | 1 ![]() |
[6] † Patriocetus kazakhstanicus Taxon name last Modified on 01/27/25 | 165 | 127 | 126 | 0 | 4 | 0 | 0 | 1 ![]() |
[7] † Orycterocetus crocodilinus Taxon name last Modified on 01/27/25 | 105 | 181 | 176 | 0 | 14 | 0 | 0 | 1 ![]() |
[8] Physeter macrocephalus Taxon name last Modified on 08/22/24 | 13 | 272 | 268 | 0 | 14 | 0 | 0 | 1 ![]() |
[9] Kogia breviceps Taxon name last Modified on 08/22/24 | 11 | 262 | 260 | 0 | 24 | 0 | 0 | 1 ![]() |
[10] Tasmacetus shepherdi Taxon name last Modified on 08/22/24 | 12 | 279 | 276 | 0 | 7 | 0 | 0 | 1 ![]() |
[11] Berardius bairdii Taxon name last Modified on 08/22/24 | 17 | 268 | 263 | 0 | 15 | 0 | 0 | 1 ![]() |
[12] Ziphius cavirostris Taxon name last Modified on 08/22/24 | 9 | 278 | 270 | 0 | 16 | 0 | 0 | 1 ![]() |
[13] Mesoplodon ginkgodens Taxon name last Modified on 08/22/24 | 29 | 250 | 249 | 0 | 17 | 0 | 0 | 1 ![]() |
[14] † Xiphiacetus bossi Taxon name last Modified on 01/27/25 | 54 | 240 | 232 | 0 | 8 | 0 | 0 | 1 ![]() |
[15] † Simocetus rayi Taxon name last Modified on 01/27/25 | 130 | 163 | 163 | 0 | 2 | 0 | 0 | 1 ![]() |
[16] † Lipotes vexillifer Taxon name last Modified on 01/27/25 | 6 | 284 | 284 | 0 | 5 | 0 | 0 | 1 ![]() |
[17] † Parapontoporia sternbergi Taxon name last Modified on 01/27/25 | 134 | 159 | 155 | 0 | 6 | 0 | 0 | 1 ![]() |
[18] Inia geoffrensis Taxon name last Modified on 08/22/24 | 5 | 290 | 277 | 0 | 13 | 0 | 0 | 1 ![]() |
[19] Pontoporia blainvillei Taxon name last Modified on 08/22/24 | 4 | 288 | 283 | 0 | 8 | 0 | 0 | 1 ![]() |
[20] † Pliopontos littoralis Taxon name last Modified on 01/27/25 | 111 | 180 | 176 | 0 | 8 | 0 | 0 | 1 ![]() |
[21] † Brachydelphis mazeasi Taxon name last Modified on 01/27/25 | 98 | 202 | 195 | 0 | 2 | 0 | 0 | 1 ![]() |
[22] † Kentriodon pernix Taxon name last Modified on 01/27/25 | 62 | 232 | 227 | 0 | 6 | 0 | 0 | 1 ![]() |
[23] † Atocetus iquensis Taxon name last Modified on 01/27/25 | 62 | 230 | 225 | 0 | 7 | 0 | 0 | 1 ![]() |
[24] † Hadrodelphis calvertense Taxon name last Modified on 01/27/25 | 122 | 166 | 165 | 0 | 8 | 0 | 0 | 1 ![]() |
[25] † Albireo whistleri Taxon name last Modified on 01/27/25 | 50 | 238 | 237 | 0 | 7 | 0 | 0 | 1 ![]() |
[26] † Denebola brachycephala Taxon name last Modified on 01/27/25 | 142 | 148 | 147 | 0 | 5 | 0 | 0 | 1 ![]() |
[27] Delphinapterus leucas Taxon name last Modified on 08/22/24 | 5 | 289 | 282 | 0 | 8 | 0 | 0 | 1 ![]() |
[28] Monodon monoceros Taxon name last Modified on 08/22/24 | 11 | 277 | 268 | 0 | 15 | 0 | 0 | 1 ![]() |
[29] † Odobenocetops peruvianus Taxon name last Modified on 01/27/25 | 107 | 162 | 162 | 0 | 26 | 0 | 0 | 1 ![]() |
[30] † Australithax intermedia Taxon name last Modified on 01/27/25 | 173 | 121 | 121 | 0 | 1 | 0 | 0 | 1 ![]() |
[31] † Lomacetus ginsburgi Taxon name last Modified on 01/27/25 | 180 | 115 | 113 | 0 | 1 | 0 | 0 | 1 ![]() |
[32] † Miophocaena nishinoi Taxon name last Modified on 01/27/25 | 209 | 86 | 86 | 0 | 0 | 0 | 0 | 1 ![]() |
[33] † Archaeophocaena teshioensis Taxon name last Modified on 01/27/25 | 238 | 55 | 55 | 0 | 2 | 0 | 0 | 1 ![]() |
[34] † Pterophocaena nishinoi Taxon name last Modified on 01/27/25 | 139 | 152 | 151 | 0 | 4 | 0 | 0 | 1 ![]() |
[35] † Haborophocoena minutus Taxon name last Modified on 01/27/25 | 185 | 110 | 109 | 0 | 1 | 0 | 0 | 1 ![]() |
[36] † Haborophocoena toyoshimai Taxon name last Modified on 01/27/25 | 120 | 174 | 174 | 0 | 1 | 0 | 0 | 1 ![]() |
[37] † Piscolithax boreios Taxon name last Modified on 01/27/25 | 137 | 153 | 152 | 0 | 5 | 0 | 0 | 1 ![]() |
[38] † Piscolithax longirostris Taxon name last Modified on 01/27/25 | 93 | 196 | 196 | 0 | 6 | 0 | 0 | 1 ![]() |
[39] † Piscolithax tedfordi Taxon name last Modified on 01/27/25 | 112 | 178 | 177 | 0 | 5 | 0 | 0 | 1 ![]() |
[40] † Salumiphocaena stocktoni Taxon name last Modified on 01/27/25 | 158 | 132 | 131 | 0 | 5 | 0 | 0 | 1 ![]() |
[41] † Septemtriocetus bosselaersi Taxon name last Modified on 01/27/25 | 202 | 86 | 86 | 0 | 7 | 0 | 0 | 1 ![]() |
[42] Phocoena sinus Taxon name last Modified on 08/22/24 | 20 | 273 | 269 | 0 | 6 | 0 | 0 | 1 ![]() |
[43] Phocoena spinipinnis Taxon name last Modified on 08/22/24 | 16 | 281 | 273 | 0 | 6 | 0 | 0 | 1 ![]() |
[44] Phocoena dioptrica Taxon name last Modified on 08/22/24 | 69 | 227 | 220 | 0 | 6 | 0 | 0 | 1 ![]() |
[45] Phocoena phocoena Taxon name last Modified on 08/22/24 | 6 | 292 | 282 | 0 | 7 | 0 | 0 | 1 ![]() |
[46] Phocoenoides dalli Taxon name last Modified on 08/22/24 | 16 | 286 | 273 | 0 | 6 | 0 | 0 | 1 ![]() |
[47] Neophocaena phocaenoides Taxon name last Modified on 08/22/24 | 11 | 287 | 277 | 0 | 7 | 0 | 0 | 1 ![]() |
[48] Lissodelphis borealis Taxon name last Modified on 08/22/24 | 23 | 273 | 266 | 0 | 6 | 0 | 0 | 1 ![]() |
[49] Cephalorhynchus hectori Taxon name last Modified on 08/22/24 | 15 | 276 | 274 | 0 | 6 | 0 | 0 | 1 ![]() |
[50] Lagenorhynchus acutus Taxon name last Modified on 08/22/24 | 29 | 260 | 259 | 0 | 7 | 0 | 0 | 1 ![]() |
[51] Orcaella brevirostris Taxon name last Modified on 08/22/24 | 36 | 255 | 251 | 0 | 8 | 0 | 0 | 1 ![]() |
[52] Orcinus orca Taxon name last Modified on 08/22/24 | 15 | 282 | 272 | 0 | 8 | 0 | 0 | 1 ![]() |
[53] Feresa attenuata Taxon name last Modified on 08/22/24 | 16 | 276 | 271 | 0 | 8 | 0 | 0 | 1 ![]() |
[54] Globicephala macrorhynchus Taxon name last Modified on 08/22/24 | 18 | 272 | 269 | 0 | 8 | 0 | 0 | 1 ![]() |
[55] Grampus griseus Taxon name last Modified on 08/22/24 | 20 | 269 | 264 | 0 | 11 | 0 | 0 | 1 ![]() |
[56] † Hemisyntrachelus cortesii Taxon name last Modified on 01/27/25 | 106 | 182 | 181 | 0 | 8 | 0 | 0 | 1 ![]() |
[57] Peponocephala electra Taxon name last Modified on 08/22/24 | 22 | 269 | 265 | 0 | 8 | 0 | 0 | 1 ![]() |
[58] Pseudorca crassidens Taxon name last Modified on 08/22/24 | 17 | 281 | 270 | 0 | 8 | 0 | 0 | 1 ![]() |
[59] Delphinus delphis Taxon name last Modified on 08/22/24 | 12 | 283 | 276 | 0 | 7 | 0 | 0 | 1 ![]() |
[60] Lagenodelphis hosei Taxon name last Modified on 08/22/24 | 18 | 276 | 270 | 0 | 7 | 0 | 0 | 1 ![]() |
[61] Sousa chinensis Taxon name last Modified on 08/22/24 | 36 | 262 | 252 | 0 | 7 | 0 | 0 | 1 ![]() |
[62] Stenella attenuata Taxon name last Modified on 08/22/24 | 15 | 284 | 273 | 0 | 7 | 0 | 0 | 1 ![]() |
[63] Tursiops truncatus Taxon name last Modified on 08/22/24 | 1 | 303 | 286 | 0 | 7 | 0 | 0 | 1 ![]() |
[64] Sotalia fluviatilis Taxon name last Modified on 08/22/24 | 20 | 274 | 268 | 0 | 7 | 0 | 0 | 1 ![]() |
[65] Steno bredanensis Taxon name last Modified on 08/22/24 | 15 | 282 | 273 | 0 | 7 | 0 | 0 | 1 ![]() |
[66] † Eodelphinus kabatensis Taxon name last Modified on 01/27/25 | 167 | 129 | 128 | 0 | 0 | 0 | 0 | 1 ![]() |
[67] † Papahu taitapu Taxon name last Modified on 01/27/25 | 154 | 139 | 138 | 0 | 3 | 0 | 0 | 1 ![]() |
[68] † Waipatia hectori Taxon name last Modified on 01/27/25 | 242 | 52 | 49 | 0 | 2 | 0 | 0 | 1 ![]() |
[69] † Awamokoa tokarahi Taxon name last Modified on 01/27/25 | 218 | 79 | 77 | 0 | 0 | 0 | 0 | 1 ![]() |
[70] † Waipatia maerewhenua Taxon name last Modified on 01/27/25 | 79 | 213 | 212 | 0 | 4 | 0 | 0 | 1 ![]() |
[71] † Otekaikea marplesi Taxon name last Modified on 01/27/25 | 127 | 168 | 165 | 0 | 1 | 0 | 0 | 1 ![]() |
[72] † Otekaikea huata Taxon name last Modified on 01/27/25 | 119 | 176 | 174 | 0 | 1 | 0 | 0 | 1 ![]() |
[73] † Squalodon calvertensis Taxon name last Modified on 01/27/25 | 64 | 230 | 215 | 0 | 3 | 0 | 0 | 1 ![]() |
[74] † Notocetus vanbenedeni Taxon name last Modified on 01/27/25 | 59 | 232 | 229 | 0 | 7 | 0 | 0 | 1 ![]() |
[75] † Squalodelphis fabianii Taxon name last Modified on 01/27/25 | 148 | 144 | 142 | 0 | 5 | 0 | 0 | 1 ![]() |
[76] † Prosqualodon davidis Taxon name last Modified on 01/27/25 | 103 | 192 | 186 | 0 | 6 | 0 | 0 | 1 ![]() |
[77] † Squaloziphius emlongi Taxon name last Modified on 01/27/25 | 165 | 125 | 124 | 0 | 6 | 0 | 0 | 1 ![]() |
[78] † Zarhachis flagellator Taxon name last Modified on 01/27/25 | 72 | 220 | 215 | 0 | 8 | 0 | 0 | 1 ![]() |
[79] † Pomatodelphis inaequalis Taxon name last Modified on 01/27/25 | 114 | 170 | 170 | 0 | 11 | 0 | 0 | 1 ![]() |
[80] † Papahu sp. zmt 73 Taxon name last Modified on 01/27/25 | 251 | 44 | 44 | 0 | 0 | 0 | 0 | 1 ![]() |
[81] Platanista gangetica Taxon name last Modified on 08/22/24 | 16 | 277 | 269 | 0 | 10 | 0 | 0 | 1 ![]() |
[82] † Phocageneus venustus Taxon name last Modified on 01/27/25 | 253 | 40 | 39 | 0 | 3 | 0 | 0 | 1 ![]() |
[83] † Phoberodon arctirostris Taxon name last Modified on 01/27/25 | 136 | 159 | 157 | 0 | 2 | 0 | 0 | 1 ![]() |
[84] † Aondelphis talen Taxon name last Modified on 01/27/25 | 229 | 64 | 64 | 0 | 2 | 0 | 0 | 1 ![]() |
[85] † Dolgopolis kinchikafiforo Taxon name last Modified on 01/27/25 | 177 | 100 | 99 | 0 | 19 | 0 | 0 | 1 ![]() |
[86] † Inticetus vertizi Taxon name last Modified on 01/27/25 | 144 | 158 | 151 | 0 | 0 | 0 | 0 | 1 ![]() |
[87] † Prosqualodon australis Taxon name last Modified on 01/27/25 | 185 | 109 | 109 | 0 | 1 | 0 | 0 | 1 ![]() |
[88] † Eosqualodon langewieschei Taxon name last Modified on 01/27/25 | 94 | 198 | 198 | 0 | 3 | 0 | 0 | 1 ![]() |
[89] † Squalodon grateloupii Taxon name last Modified on 01/27/25 | 65 | 227 | 226 | 0 | 3 | 0 | 0 | 1 ![]() |
[90] † Allodelphis pratti Taxon name last Modified on 01/27/25 | 161 | 131 | 131 | 0 | 3 | 0 | 0 | 1 ![]() |
[91] † Arktocara yakataga Taxon name last Modified on 01/27/25 | 195 | 96 | 96 | 0 | 4 | 0 | 0 | 1 ![]() |
[92] † Ninjadelphis ujiharai Taxon name last Modified on 01/27/25 | 163 | 129 | 128 | 0 | 4 | 0 | 0 | 1 ![]() |
[93] † Zarhinocetus errabundus Taxon name last Modified on 01/27/25 | 115 | 175 | 175 | 0 | 5 | 0 | 0 | 1 ![]() |
[94] † Goedertius oregonensis Taxon name last Modified on 01/27/25 | 161 | 127 | 127 | 0 | 7 | 0 | 0 | 1 ![]() |
[95] † Huaridelphis raimondii Taxon name last Modified on 01/27/25 | 119 | 175 | 173 | 0 | 3 | 0 | 0 | 1 ![]() |
[96] † Medocinia tetragorhina Taxon name last Modified on 01/27/25 | 208 | 84 | 84 | 0 | 3 | 0 | 0 | 1 ![]() |
[97] † Macrosqualodelphis ukupachai Taxon name last Modified on 01/27/25 | 150 | 141 | 139 | 0 | 6 | 0 | 0 | 1 ![]() |
[98] † Dilophodelphis fordycei Taxon name last Modified on 01/27/25 | 168 | 123 | 123 | 0 | 4 | 0 | 0 | 1 ![]() |
[99] † Araeodelphis natator Taxon name last Modified on 01/27/25 | 206 | 87 | 86 | 0 | 3 | 0 | 0 | 1 ![]() |
[100] † Squalodon whitmorei Taxon name last Modified on 01/27/25 | 160 | 133 | 132 | 0 | 3 | 0 | 0 | 1 ![]() |
[101] † Squalodon murdochi Taxon name last Modified on 01/27/25 | 230 | 65 | 65 | 0 | 0 | 0 | 0 | 1 ![]() |
[102] † Squalodon bellunensis Taxon name last Modified on 01/27/25 | 80 | 213 | 211 | 0 | 3 | 0 | 0 | 1 ![]() |
[103] † Squalodon peregrinus Taxon name last Modified on 01/27/25 | 208 | 85 | 85 | 0 | 2 | 0 | 0 | 1 ![]() |
[104] † Eosqualodon latirostris Taxon name last Modified on 01/27/25 | 192 | 100 | 100 | 0 | 3 | 0 | 0 | 1 ![]() |
Project downloads 
type | number of downloads | Individual items downloaded (where applicable) |
Total downloads from project | 2 | |
Matrix downloads | 1 | Phylogenetic matrix (1 download); |
Project downloads | 1 |