Project 2044: Fordyce, R.E., Marx, F.G. 2018. Gigantism precedes filter feeding in baleen whale evolution. Current Biology. 28:1670-1676.e2.
Abstract
Baleen whales (Mysticeti) are the largest animals on Earth, thanks to their ability to filter feed huge volumes of small prey from seawater. Mysticetes appeared during the Late Eocene, but evidence of their early evolution remains both sparse and controversial, with several models competing to explain the origin of baleen-based bulk feeding. Here, we describe a virtually complete skull of Llanocetus denticrenatus, the second-oldest (ca 34 Ma) mysticete known. The new material represents the same individual as the type and only specimen, a fragmentary mandible. Phylogenetic analysis groups Llanocetus with the oldest mysticete, Mystacodon selenensis, into the basal family Llanocetidae. Llanocetus is gigantic (body length ca 8 m) compared to other early mysticetes. The broad rostrum has sharp, widely-spaced teeth with marked dental abrasion and attrition suggesting biting and occlusal shearing. As in extant mysticetes, the palate bears many sulci, commonly interpreted as osteological correlates of baleen. Unexpectedly, these sulci converge on the upper alveoli, suggesting a peri-dental blood supply to well-developed gums, rather than to inter-alveolar racks of baleen. We interpret Llanocetus as a raptorial or suction feeder, revealing that whales evolved gigantism well before the emergence of filter feeding. Rather than driving the origin of mysticetes, baleen and filtering likely only arose following an initial phase of suction-assisted raptorial feeding. This scenario strikingly differs from that proposed for odontocetes, whose defining adaptation ? echolocation ? was present even in their earliest representatives.Read the article »
Article DOI: 10.1016/j.cub.2018.04.027
Project DOI: 10.7934/P2044, http://dx.doi.org/10.7934/P2044
This project contains | Matrices |
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Download Project SDD File ![]() | Total scored cells: 29213 Total media associated with cells: 17480 Total labels associated with cell media: 18229 |
Characters | |
Total characters: 275 Total characters with associated media: 0 Total characters with media with labels: 0 Total character states: 645 Total character states with associated media: 0 Total character states with media with labels:0 Total unordered/ordered characters:237/38 |
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MorphoBank Project 2044
MorphoBank Project 2044
- Creation Date:
23 May 2015 - Publication Date:
05 June 2018 - Project views: 123208
- Media downloads: 51
- Matrix downloads: 74
Authors' Institutions
- Museum Victoria, Melbourne
- National Museum of Natural History, Smithsonian Institution
- Monash University
- University of Otago
- Royal Belgian Institute of Natural Sciences
- Smithsonian Institution
Members
member name | taxa ![]() |
specimens ![]() |
media ![]() | media notes | chars ![]() | character
| cell scorings (scored, NPA, "-") | cell
| rules ![]() | ||||||||||
Felix Marx Project Administrator | 106 | 294 | 5617 | 53 | 275 | 219 | 0 | 0 | 29213 (16140, 11841, 1232) | 37 | 14849 | 18229 | 0 | ||||||
Seth Kaufman | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | 0 |
Taxonomic Overview for Matrix 'M3690' (106 Taxa)
taxon | unscored cells |
scored cells ![]() |
no cell support ![]() |
NPA cells |
"-" cells | cell images | labels on cell images |
member access |
[1] † Zygorhiza kochii Last Modified in 06/17/15 | 0 | 250 | 0 | 18 | 7 | 313 | 332 | 2 ![]() |
[2] † Albertocetus meffordorum Last Modified in 12/04/17 | 0 | 188 | 0 | 75 | 12 | 275 | 280 | 2 ![]() |
[3] † Archaeodelphis patrius Last Modified in 06/16/15 | 0 | 102 | 0 | 164 | 9 | 119 | 126 | 2 ![]() |
[4] † Olympicetus avitus Last Modified in 11/07/17 | 0 | 137 | 0 | 129 | 9 | 152 | 159 | 2 ![]() |
[5] † Waipatia maerewhenua Last Modified in 06/17/15 | 0 | 217 | 0 | 45 | 13 | 247 | 261 | 2 ![]() |
[6] Physeter macrocephalus Last Modified in 12/15/17 | 0 | 227 | 2 | 11 | 39 | 263 | 276 | 2 ![]() |
[7] † Aetiocetus cotylalveus Last Modified in 06/16/15 | 0 | 164 | 0 | 109 | 2 | 197 | 212 | 2 ![]() |
[8] † Aetiocetus polydentatus Last Modified in 06/16/15 | 0 | 175 | 22 | 97 | 3 | 192 | 206 | 2 ![]() |
[9] † Aetiocetus weltoni Last Modified in 06/16/15 | 0 | 188 | 0 | 84 | 3 | 220 | 230 | 2 ![]() |
[10] † Aglaocetus moreni Last Modified in 06/16/15 | 0 | 140 | 0 | 123 | 12 | 156 | 158 | 2 ![]() |
[11] † Aglaocetus patulus Last Modified in 06/16/15 | 0 | 187 | 0 | 77 | 13 | 224 | 235 | 2 ![]() |
[12] † Archaebalaenoptera castriarquati Last Modified in 06/16/15 | 0 | 98 | 0 | 160 | 17 | 119 | 122 | 2 ![]() |
[13] † Aulocetus latus Last Modified in 12/15/17 | 0 | 99 | 0 | 170 | 6 | 113 | 118 | 2 ![]() |
[14] † Balaena montalionis Last Modified in 06/16/15 | 0 | 71 | 0 | 186 | 18 | 86 | 90 | 2 ![]() |
[15] Balaena mysticetus Last Modified in 12/05/13 | 0 | 253 | 0 | 2 | 23 | 309 | 317 | 2 ![]() |
[16] † Balaena ricei Last Modified in 06/16/15 | 0 | 96 | 0 | 161 | 18 | 116 | 122 | 2 ![]() |
[17] † Balaenella brachyrhynus Last Modified in 06/16/15 | 0 | 134 | 0 | 121 | 20 | 171 | 175 | 2 ![]() |
[18] Balaenoptera acutorostrata Last Modified in 12/05/13 | 0 | 259 | 0 | 0 | 17 | 284 | 301 | 2 ![]() |
[19] † Balaenoptera bertae Last Modified in 06/16/15 | 0 | 70 | 0 | 198 | 7 | 94 | 99 | 2 ![]() |
[20] Balaenoptera bonaerensis Last Modified in 12/05/13 | 0 | 254 | 0 | 5 | 16 | 299 | 312 | 2 ![]() |
[21] Balaenoptera borealis Last Modified in 12/05/13 | 0 | 258 | 0 | 1 | 17 | 279 | 298 | 2 ![]() |
[22] Balaenoptera musculus Last Modified in 12/05/13 | 0 | 256 | 0 | 3 | 17 | 295 | 310 | 2 ![]() |
[23] Balaenoptera omurai Last Modified in 12/05/13 | 0 | 252 | 246 | 7 | 16 | 0 | 0 | 2 ![]() |
[24] Balaenoptera physalus Last Modified in 12/05/13 | 0 | 259 | 0 | 1 | 17 | 291 | 310 | 2 ![]() |
[25] † Balaenoptera portisi Last Modified in 06/16/15 | 0 | 180 | 0 | 83 | 12 | 227 | 238 | 2 ![]() |
[26] † Balaenoptera ryani Last Modified in 06/17/15 | 0 | 81 | 79 | 190 | 5 | 1 | 1 | 2 ![]() |
[27] † Balaenoptera siberi Last Modified in 06/17/15 | 0 | 158 | 0 | 103 | 16 | 184 | 190 | 2 ![]() |
[28] † Balaenula astensis Last Modified in 06/17/15 | 0 | 146 | 0 | 110 | 19 | 188 | 188 | 2 ![]() |
[29] † Balaenula sp. Last Modified in 06/17/15 | 0 | 125 | 0 | 132 | 18 | 149 | 154 | 2 ![]() |
[30] † Brandtocetus chongulek Last Modified in 06/17/15 | 0 | 122 | 0 | 148 | 6 | 151 | 165 | 2 ![]() |
[31] Caperea marginata Last Modified in 12/05/13 | 0 | 252 | 0 | 3 | 26 | 359 | 376 | 2 ![]() |
[32] † Cephalotropis coronatus Last Modified in 06/17/15 | 0 | 138 | 136 | 125 | 12 | 1 | 1 | 2 ![]() |
[33] † Cetotherium megalophysum Last Modified in 12/15/17 | 0 | 117 | 114 | 142 | 17 | 0 | 0 | 2 ![]() |
[34] † Cetotherium rathkii Last Modified in 06/17/15 | 0 | 139 | 0 | 118 | 18 | 165 | 170 | 2 ![]() |
[35] † Cetotherium riabinini Last Modified in 06/17/15 | 0 | 185 | 0 | 73 | 17 | 215 | 220 | 2 ![]() |
[36] † Chonecetus sookensis Last Modified in 06/17/15 | 0 | 79 | 0 | 194 | 2 | 93 | 99 | 2 ![]() |
[37] † Coronodon havensteini Last Modified in 08/25/17 | 0 | 201 | 0 | 70 | 5 | 269 | 277 | 2 ![]() |
[38] † Diorocetus chichibuensis Last Modified in 06/17/15 | 0 | 176 | 0 | 86 | 13 | 210 | 213 | 2 ![]() |
[39] † Diorocetus hiatus Last Modified in 06/17/15 | 0 | 213 | 0 | 49 | 13 | 263 | 279 | 2 ![]() |
[40] † Diorocetus shobarensis Last Modified in 06/17/15 | 0 | 127 | 0 | 136 | 13 | 160 | 164 | 2 ![]() |
[41] † Diunatans luctoretemergo Last Modified in 06/17/15 | 0 | 123 | 0 | 149 | 4 | 149 | 159 | 2 ![]() |
[42] † Eomysticetus whitmorei Last Modified in 06/17/15 | 0 | 181 | 0 | 79 | 15 | 214 | 221 | 2 ![]() |
[43] † Eschrichtioides gastaldii Last Modified in 06/17/15 | 0 | 144 | 0 | 113 | 18 | 170 | 174 | 2 ![]() |
[44] Eschrichtius robustus Last Modified in 12/05/13 | 0 | 263 | 0 | 0 | 17 | 320 | 334 | 2 ![]() |
[45] † Eubalaena ianitrix Last Modified in 11/07/17 | 0 | 69 | 0 | 200 | 6 | 93 | 95 | 2 ![]() |
[46] † Eubalaena shinshuensis Last Modified in 06/17/15 | 0 | 55 | 0 | 202 | 18 | 77 | 77 | 2 ![]() |
[47] Eubalaena spp. Last Modified in 12/05/13 | 0 | 258 | 0 | 0 | 23 | 403 | 418 | 2 ![]() |
[48] † Fragilicetus velponi Last Modified in 11/02/16 | 0 | 87 | 1 | 185 | 4 | 99 | 107 | 2 ![]() |
[49] † Fucaia buelli Last Modified in 08/03/15 | 0 | 148 | 0 | 124 | 3 | 175 | 183 | 2 ![]() |
[50] † Fucaia goedertorum Last Modified in 12/16/15 | 0 | 192 | 0 | 79 | 4 | 235 | 240 | 2 ![]() |
[51] † Gricetoides aurorae Last Modified in 06/17/15 | 0 | 82 | 0 | 192 | 1 | 96 | 105 | 2 ![]() |
[52] † Herentalia nigra Last Modified in 02/08/16 | 0 | 67 | 8 | 201 | 7 | 72 | 77 | 2 ![]() |
[53] † Herpetocetus bramblei Last Modified in 06/17/15 | 0 | 145 | 0 | 113 | 17 | 181 | 199 | 2 ![]() |
[54] † Herpetocetus morrowi Last Modified in 02/17/16 | 0 | 217 | 0 | 41 | 19 | 329 | 347 | 2 ![]() |
[55] † Herpetocetus sp. Last Modified in 06/17/15 | 0 | 64 | 0 | 204 | 7 | 78 | 81 | 2 ![]() |
[56] † Herpetocetus transatlanticus Last Modified in 06/17/15 | 0 | 116 | 0 | 152 | 7 | 153 | 171 | 2 ![]() |
[57] † Horopeta umarere Last Modified in 03/29/16 | 0 | 118 | 0 | 145 | 12 | 139 | 140 | 2 ![]() |
[58] † Incakujira anillodefuego Last Modified in 10/02/16 | 0 | 203 | 3 | 58 | 16 | 245 | 249 | 2 ![]() |
[59] † Isanacetus laticephalus Last Modified in 06/17/15 | 0 | 197 | 0 | 64 | 15 | 244 | 251 | 2 ![]() |
[60] † Janjucetus hunderi Last Modified in 06/17/15 | 0 | 181 | 0 | 88 | 6 | 191 | 201 | 2 ![]() |
[61] † Joumocetus shimizui Last Modified in 06/17/15 | 0 | 116 | 0 | 144 | 15 | 136 | 141 | 2 ![]() |
[62] † Kurdalagonus mchedlidzei Last Modified in 06/17/15 | 0 | 129 | 0 | 130 | 16 | 157 | 167 | 2 ![]() |
[63] † Llanocetus denticrenatus Last Modified in 06/17/15 | 0 | 179 | 0 | 91 | 5 | 225 | 232 | 2 ![]() |
[64] † Mammalodon colliveri Last Modified in 06/17/15 | 0 | 175 | 0 | 93 | 7 | 206 | 219 | 2 ![]() |
[65] † Mammalodon hakataramea Last Modified in 09/11/15 | 0 | 37 | 0 | 239 | 0 | 41 | 39 | 2 ![]() |
[66] † Matapanui waihao Last Modified in 08/25/17 | 0 | 121 | 0 | 139 | 15 | 147 | 153 | 2 ![]() |
[67] † Mauicetus parki Last Modified in 06/17/15 | 0 | 183 | 147 | 80 | 13 | 37 | 40 | 2 ![]() |
[68] † Megaptera hubachi Last Modified in 12/15/17 | 0 | 183 | 0 | 75 | 17 | 220 | 228 | 2 ![]() |
[69] † Megaptera miocaena Last Modified in 12/15/17 | 0 | 150 | 0 | 107 | 18 | 192 | 209 | 2 ![]() |
[70] Megaptera novaeangliae Last Modified in 12/05/13 | 0 | 261 | 0 | 1 | 15 | 329 | 346 | 2 ![]() |
[71] † Metopocetus durinasus Last Modified in 06/17/15 | 0 | 81 | 0 | 188 | 6 | 103 | 114 | 2 ![]() |
[72] † Metopocetus hunteri Last Modified in 03/29/16 | 0 | 107 | 0 | 162 | 7 | 132 | 144 | 2 ![]() |
[73] † Metopocetus vandelli Last Modified in 12/15/17 | 0 | 101 | 0 | 169 | 6 | 122 | 129 | 2 ![]() |
[74] † Micromysticetus rothauseni Last Modified in 06/17/15 | 0 | 90 | 0 | 183 | 2 | 105 | 114 | 2 ![]() |
[75] † Miocaperea pulchra Last Modified in 06/17/15 | 0 | 137 | 0 | 118 | 21 | 165 | 173 | 2 ![]() |
[76] † Morawanocetus yabukii Last Modified in 06/17/15 | 0 | 140 | 64 | 130 | 5 | 92 | 94 | 2 ![]() |
[77] † Morenocetus parvus Last Modified in 06/17/15 | 0 | 113 | 0 | 157 | 5 | 144 | 150 | 2 ![]() |
[78] † Mystacodon selenensis Last Modified in 08/25/17 | 0 | 127 | 9 | 143 | 5 | 135 | 139 | 2 ![]() |
[79] † Nannocetus eremus Last Modified in 06/17/15 | 0 | 89 | 0 | 184 | 2 | 113 | 123 | 2 ![]() |
[80] † NMNS PV19540 Last Modified in 12/29/16 | 0 | 168 | 166 | 90 | 18 | 0 | 0 | 2 ![]() |
[81] † NMNZ MM001630 Last Modified in 06/17/15 | 0 | 195 | 183 | 62 | 18 | 0 | 0 | 2 ![]() |
[82] † OCPC 1178 Last Modified in 06/17/15 | 0 | 91 | 89 | 182 | 2 | 0 | 0 | 2 ![]() |
[83] † OU 22224 Last Modified in 06/17/15 | 0 | 107 | 104 | 166 | 2 | 0 | 0 | 2 ![]() |
[84] † OU 22705 Last Modified in 06/17/15 | 0 | 151 | 149 | 110 | 14 | 0 | 0 | 2 ![]() |
[85] † OU GS10897 Last Modified in 06/17/15 | 0 | 56 | 55 | 217 | 3 | 0 | 0 | 2 ![]() |
[86] † Parabalaenoptera baulinensis Last Modified in 06/17/15 | 0 | 162 | 0 | 98 | 15 | 190 | 193 | 2 ![]() |
[87] † Parietobalaena campiniana Last Modified in 06/17/15 | 0 | 153 | 0 | 108 | 14 | 191 | 193 | 2 ![]() |
[88] † Parietobalaena palmeri Last Modified in 06/17/15 | 0 | 217 | 0 | 50 | 13 | 280 | 289 | 2 ![]() |
[89] † Parietobalaena sp. Last Modified in 06/17/15 | 0 | 154 | 0 | 106 | 15 | 179 | 188 | 2 ![]() |
[90] † Parietobalaena yamaokai Last Modified in 06/17/15 | 0 | 174 | 0 | 91 | 13 | 212 | 218 | 2 ![]() |
[91] † Pelocetus calvertensis Last Modified in 06/17/15 | 0 | 220 | 0 | 41 | 14 | 252 | 255 | 2 ![]() |
[92] † Peripolocetus vexillifer Last Modified in 06/17/15 | 0 | 60 | 58 | 214 | 1 | 1 | 1 | 2 ![]() |
[93] † Pinocetus polonicus Last Modified in 06/17/15 | 0 | 115 | 0 | 148 | 12 | 123 | 125 | 2 ![]() |
[94] † Piscobalaena nana Last Modified in 06/17/15 | 0 | 242 | 0 | 15 | 19 | 300 | 309 | 2 ![]() |
[95] † Plesiobalaenoptera quarantellii Last Modified in 06/17/15 | 0 | 137 | 0 | 121 | 17 | 148 | 157 | 2 ![]() |
[96] † Thinocetus arthritus Last Modified in 06/17/15 | 0 | 143 | 0 | 120 | 12 | 170 | 176 | 2 ![]() |
[97] † Tiphyocetus temblorensis Last Modified in 06/17/15 | 0 | 71 | 0 | 204 | 0 | 81 | 84 | 2 ![]() |
[98] † Titanocetus sammarinensis Last Modified in 06/17/15 | 0 | 139 | 0 | 125 | 11 | 156 | 160 | 2 ![]() |
[99] † Tiucetus rosae Last Modified in 09/15/17 | 0 | 139 | 0 | 135 | 1 | 173 | 177 | 2 ![]() |
[100] † Tohoraata raekohao Last Modified in 03/31/16 | 0 | 113 | 0 | 160 | 2 | 126 | 133 | 2 ![]() |
[101] † Tokarahia kauaeroa Last Modified in 03/31/16 | 0 | 149 | 0 | 113 | 13 | 172 | 176 | 2 ![]() |
[102] † Uranocetus gramensis Last Modified in 06/17/15 | 0 | 173 | 0 | 86 | 16 | 191 | 196 | 2 ![]() |
[103] † Waharoa ruwhenua Last Modified in 11/07/16 | 0 | 198 | 0 | 62 | 17 | 309 | 313 | 2 ![]() |
[104] † Whakakai waipata Last Modified in 01/26/17 | 0 | 80 | 0 | 194 | 1 | 95 | 99 | 2 ![]() |
[105] † Yamatocetus canaliculatus Last Modified in 06/17/15 | 0 | 191 | 0 | 70 | 14 | 218 | 220 | 2 ![]() |
[106] † ZMT 67 Last Modified in 06/17/15 | 0 | 140 | 139 | 122 | 13 | 0 | 0 | 2 ![]() |
Project views 
type | number of views | Individual items viewed (where applicable) |
Total project views | 123208 | |
Project overview | 2280 | |
Matrix views | 1462 | Matrix landing page (999 views); Mysticetes mark II (463 views); |
Documents list | 616 | |
Media views | 41846 | Media search (2595 views); M462871 (1 view); M344531 (2 views); M344454 (3 views); M344461 (8 views); M344446 (1 view); M344584 (1 view); M410639 (4 views); M340860 (206 views); M340865 (220 views); M340809 (274 views); M340871 (194 views); M340858 (190 views); M340804 (275 views); M340881 (229 views); M340856 (199 views); M340837 (213 views); M340819 (248 views); M340799 (246 views); M340788 (277 views); M340816 (274 views); M340872 (206 views); M340829 (225 views); M340854 (197 views); M340863 (204 views); M340883 (196 views); M340813 (262 views); M340847 (195 views); M340840 (195 views); M340796 (291 views); M340857 (238 views); M340817 (257 views); M340807 (281 views); M340874 (219 views); M340827 (265 views); M340823 (236 views); M340879 (206 views); M340794 (296 views); M340802 (297 views); M340844 (203 views); M340864 (211 views); M340834 (210 views); M340835 (295 views); M340831 (220 views); M340885 (205 views); M340789 (286 views); M340848 (201 views); M340797 (251 views); 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Views for media list | 2794 | |
Specimen list | 59710 | |
Taxon list | 7669 | |
Bibliography | 6831 |
Project downloads 
type | number of downloads | Individual items downloaded (where applicable) |
Total downloads from project | 297 | |
Matrix downloads | 74 | Mysticetes mark II (74 downloads); |
Document downloads | 12 | Palatal sulci and body size optimisation (4 downloads); Institutional abbreviations (1 download); Supplementary Nexus file (7 downloads); |
Project downloads | 160 | |
Media downloads | 51 | M379508 (1 download); M379497 (2 downloads); M379494 (1 download); M379495 (2 downloads); M379496 (2 downloads); M379501 (1 download); M379500 (1 download); M379499 (1 download); M379503 (2 downloads); M379510 (2 downloads); M410674 (2 downloads); M343097 (1 download); M343098 (1 download); M343099 (1 download); M343104 (1 download); M343105 (1 download); M343111 (1 download); M343209 (1 download); M343211 (1 download); M342550 (1 download); M342553 (1 download); M342556 (1 download); M342570 (1 download); M342559 (1 download); M342561 (1 download); M342565 (1 download); M342568 (1 download); M342571 (1 download); M342581 (1 download); M344011 (1 download); M381629 (1 download); M379498 (1 download); M340788 (1 download); M340790 (1 download); M340832 (1 download); M343531 (1 download); M345671 (1 download); M343486 (1 download); M343487 (1 download); M343489 (1 download); M343493 (1 download); M343494 (1 download); M343495 (1 download); M343526 (1 download); M343501 (1 download); |