Pseudotoothed Birds (Aves, Odontopterygiformes) from the Early Tertiary of Morocco Author Bourdon, Estelle Author Amaghzaz, Mbarek Author Bouya, Baâdi text American Museum Novitates 2010 2010-12-20 2010 3704 1 72 http://www.bioone.org/doi/abs/10.1206/3704.2 journal article 5893 10.1206/3704.2 1577e5f5-b4ef-4ba8-a7c2-d9a47fa0b6a0 0003-0082 4597660 Dasornis toliapica ( Owen, 1873 ) , new combination Odontopteryx toliapica Owen, 1873 . Macrodontopteryx oweni Harrison and Walker, 1976 . Neptuniavis minor Harrison and Walker, 1977 . REFERRED MATERIAL: OCP.DEK/GE 534B, distal part of right tibiotarsus; OCP.DEK/GE 1001, fragments of cranium; OCP.DEK/GE 1002, proximal part of right ulna; OCP.DEK/GE 1003, incomplete proximal part of leπ humerus; OCP.DEK/GE 1005, proximal part of leπ ulna; OCP.DEK/GE 1011, distal part of leπ tibiotarsus; OCP.DEK/GE 1012, proximal part of leπ carpometacarpus; OCP.DEK/GE 1013, fragment of shaπ of humerus + fragment of shaπ of ulna + distal part of leπ tibiotarsus; OCP.DEK/GE 1017, distal part of leπ tibiotarsus; OCP.DEK/GE 1018, distal part of right ulna; OCP.DEK/GE 1020, incomplete distal end of right tarsometatarsus; OCP.DEK/GE 1024, fragment of shaπ of leπ carpometacarpus; OCP.DEK/GE 1026, fragment of shaπ of humerus; OCP.DEK/GE 1029, fragment of distal part of right tarsometatarsus; OCP.DEK/GE 1034, distal part of leπ ulna; OCP.DEK/GE 1035, distal part of right ulna; OCP. DEK/GE 1038, distal part of right carpometacarpus; OCP.DEK/GE 1039, incomplete proximal part of leπ ulna; OCP.DEK/GE 1042, incomplete cranium; OCP.DEK/GE 1043, incomplete cranium; OCP.DEK/GE 1044, incomplete cranium; OCP.DEK/GE 1045, shaπ of right femur; OCP.DEK/GE 1048, proximal part of leπ ulna; OCP.DEK/GE 1051, fragment of shaπ of ulna; OCP.DEK/GE 1055, fragment of shaπ of leπ radius; OCP.DEK/GE 1056, fragment of shaπ of right radius; OCP.DEK/GE 1058, distal end of right radius; OCP.DEK/GE 1068, fragment of shaπ of right radius close to distal end; OCP.DEK/GE 1070, nearly complete right humerus; OCP.DEK/GE 1071, distal part of right ulna; OCP.DEK/GE 1073, fragments of right humerus; OCP.DEK/GE 1075, distal end of right humerus; OCP.DEK/GE 1076, incomplete cranium; OCP.DEK/GE 1079, distal part of leπ ulna lacking extremity; OCP.DEK/GE 1080, fragment of shaπ of leπ ulna; OCP.DEK/GE 1081, fragment of shaπ of leπ ulna; OCP.DEK/GE 1082, fragment of leπ carpometacarpus; OCP.DEK/GE 1083, proximal part of leπ ulna; OCP.DEK/GE 1086, proximal part of leπ humerus; OCP.DEK/GE 1088, three fragments of right coracoideum; OCP.DEK/GE 1090, fragments of distal part of leπ tibiotarsus; OCP.DEK/GE 1091, fragments of distal part of leπ tibiotarsus; OCP.DEK/GE 1092, fragment of shaπ of radius; OCP.DEK/GE 1100, fragment of shaπ of leπ carpometacarpus; OCP.DEK/GE 1101, distal part of right carpometacarpus; OCP.DEK/GE 1102, distal part of right carpometacarpus; OCP.DEK/GE 1105, proximal part of right humerus; OCP.DEK/GE 1107, fragment of shaπ and incomplete distal part of right humerus; OCP.DEK/GE 1108, fragment of shaπ of right ulna; OCP.DEK/GE 1110, distal part of leπ ulna; OCP.DEK/GE 1112, fragment of major metacarpal of right carpometacarpus; OCP.DEK/GE 1114, fragments of shaπ of leπ ulna; OCP.DEK/GE 1115, proximal part of right humerus; OCP.DEK/GE 1116, fragment of shaπ of leπ humerus in proximal half; OCP.DEK/GE 1118, distal end of right radius; OCP.DEK/GE 1121, fragment of shaπ of radius; OCP.DEK/GE 1122, fragment of shaπ of carpometacarpus; OCP.DEK/GE 1124, fragment of shaπ of leπ humerus; OCP.DEK/GE 1125, distal part of right ulna; OCP.DEK/GE 1126, proximal end of right ulna; OCP.DEK/GE 1128, fragment of shaπ of right humerus; OCP.DEK/GE 1135, fragment of shaπ of leπ ulna; OCP.DEK/GE 1137, fragment of leπ carpometacarpus; OCP.DEK/GE 1138, fragment of shaπ of radius + distal part of right carpometacarpus; OCP. DEK/GE 1139, fragment of leπ carpometacarpus; OCP.DEK/GE 1140, distal end of leπ tibiotarsus; OCP.DEK/GE 1141, fragment of shaπ of right humerus; OCP.DEK/GE 1142, fragment of shaπ of leπ ulna; OCP.DEK/GE 1144, fragment of distal end of right femur; OCP. DEK/GE 1146, distal end of leπ tarsometatarsus; OCP.DEK/GE 1147, distal end of right ulna; OCP.DEK/GE 1148, fragment of shaπ of right ulna; OCP.DEK/GE 1149, fragment of shaπ of leπ ulna; OCP.DEK/GE 1150, fragment of shaπ of leπ ulna close to proximal end; OCP.DEK/ GE 1152, proximal end of leπ carpometacarpus; OCP.DEK/GE 1153, fragment of shaπ of leπ ulna + fragment of shaπ of radius; OCP.DEK/GE 1155, fragment of shaπ of leπ tarsometatarsus close to distal end; OCP.DEK/GE 1157, fragment of shaπ of right radius close to distal end; OCP.DEK/GE 1160, fragment of shaπ of right ulna; OCP.DEK/GE 1161, fragment of shaπ of leπ ulna; OCP.DEK/GE 1163, fragment of shaπ of leπ ulna; OCP.DEK/GE 1164, several fragments of right humerus; OCP.DEK/GE 1166, portion of right mandible; OCP.DEK/ GE 1168, fragment of shaπ of humerus; OCP.DEK/GE 1171, incomplete distal part of leπ tibiotarsus; OCP.DEK/GE 1172, leπ carpometacarpus with incomplete extremities; OCP.DEK/ GE 1173A, proximal part of leπ humerus with incomplete end; OCP.DEK/GE 1174, fragment of shaπ of right tibiotarsus; OCP.DEK/GE 1175, proximal part of leπ radius with incomplete end; OCP.DEK/GE 1176, nearly complete right tarsometatarsus; OCP.DEK/GE 1179, proximal part of right humerus lacking extremity; OCP.DEK/GE 1180, fragment of right humerus close to proximal end; OCP.DEK/GE 1182, incomplete distal end of right tarsometatarsus; OCP. DEK/GE 1183, incomplete proximal part of right humerus + shaπ of ulna; OCP.DEK/GE 1185, portion of maxilla close to proximal end; OCP.DEK/GE 1187, fragments of shaπ of leπ humerus close to proximal end; OCP.DEK/GE 1188, fragments of proximal part of right humerus; OCP.DEK/GE 1190, shaπ of right tarsometatarsus; OCP.DEK/GE 1192, distal end of leπ carpometacarpus; OCP.DEK/GE 1193, distal end of right ulna; OCP.DEK/GE 1194, proximal part of right humerus; OCP.DEK/GE 1195, distal part of right humerus and proximal part of leπ coracoideum; OCP.DEK/GE 1197, proximal part of right humerus; OCP.DEK/ GE 1198, distal part of right ulna; OCP.DEK/GE 1199, fragment of shaπ of right radius close to distal end; OCP.DEK/GE 1203, incomplete distal end of leπ humerus; OCP.DEK/GE 1204, distal part of right tibiotarsus lacking extremity; OCP.DEK/GE 1205, fragment of shaπ of right humerus close to distal end; OCP.DEK/GE 1206, distal part of leπ carpometacarpus; OCP. DEK/GE 1207, fragment of shaπ of right ulna; OCP.DEK/GE 1209, several fragments of incomplete leπ humerus; OCP.DEK/GE 1210, fragment of shaπ of leπ humerus; OCP.DEK/ GE 1212, proximal part of leπ radius; OCP.DEK/GE 1213, fragment of shaπ of radius; OCP. DEK/GE 1216, proximal end of leπ humerus; OCP.DEK/GE 1217, fragment of shaπ of right femur; OCP.DEK/GE 1218, distal part of leπ humerus with incomplete end; OCP.DEK/GE 1219, fragment of shaπ of ulna; OCP.DEK/GE 1222, fragment of shaπ of right humerus; OCP. DEK/GE 1223, fragment of shaπ of leπ ulna; OCP.DEK/GE 1227, fragment of shaπ of leπ tibiotarsus; OCP.DEK/GE 1228, fragment of shaπ of leπ humerus; OCP.DEK/GE 1229, proximal end of leπ humerus; OCP.DEK/GE 1230, fragment of cranium and fragment of maxilla embedded in matrix; OCP.DEK/GE 1233, complete right humerus embedded in matrix; OCP. DEK/GE 1235, distal end of right tarsometatarsus embedded in matrix; OCP.DEK/GE 1238, cranium embedded in matrix; OCP.DEK/GE 1240, complete right humerus embedded in matrix; OCP.DEK/GE 1241, complete right carpometacarpus embedded in matrix; OCP.DEK/ GE 1243, fragment of shaπ of right ulna embedded in matrix; OCP.DEK/GE 1244, fragment of shaπ of ulna; OCP.DEK/GE 1246, distal end of leπ humerus + fragment of shaπ of ulna; OCP.DEK/GE 1250, fragment of shaπ of ulna; OCP.DEK/GE 1255, distal part of right carpometacarpus; OCP.DEK/GE 1256, distal part of leπ tibiotarsus; OCP.DEK/GE 1257, fragment of shaπ of leπ ulna; OCP.DEK/GE 1258, fragment of shaπ of right humerus; OCP.DEK/GE 1260, fragment of shaπ of ulna; OCP.DEK/GE 1262, fragment of shaπ of right ulna; OCP. DEK/GE 1264, fragment of shaπ of humerus; OCP.DEK/GE 1265, fragment of shaπ of leπ radius; OCP.DEK/GE 1270, fragment of shaπ of right humerus; OCP.DEK/GE 1272, distal end of leπ tarsometatarsus; OCP.DEK/GE 1274, fragment of shaπ of humerus; OCP.DEK/GE 1275, distal end of leπ ulna; OCP.DEK/GE 1283, proximal part of leπ ulna; OCP.DEK/GE 1287, cranium embedded in matrix; OCP.DEK/GE 1289, complete leπ humerus embedded in matrix; OCP.DEK/GE 1293, distal end of leπ tarsometatarsus; OCP.DEK/GE 1294, distal end of leπ tarsometatarsus; OCP.DEK/GE 1298, proximal part of right radius; OCP.DEK/GE 1299, distal part of right ulna lacking extremity; OCP.DEK/GE 1314, fragment of shaπ of humerus; OCP.DEK/GE 1316, incomplete proximal part of right humerus; OCP.DEK/GE 1324, incomplete proximal part of right humerus; OCP.DEK/GE 1326, proximal part of leπ radius; OCP. DEK/GE 1327, distal part of right radius; MHNL 20-149211, distal part of right humerus; MHNL 20-149212, shaπ of leπ humerus; MHNL 20-149213, fragment of shaπ of leπ humerus; MHNL 20-149214, distal part of right humerus; MHNL 20-149215, leπ ulna + fragment of shaπ of radius; MHNL 20-149216, fragments of shaπ of leπ humerus with incomplete distal end; MHNL 20-149217, fragment of shaπ of leπ ulna close to distal end; MHNL 20-149218, distal part of right ulna; MHNL 20-149219, fragment of shaπ of leπ radius in proximal part; MHNL 20-149220, two fragments of distal part of leπ ulna; MHNL 20-149221, fragment of shaπ of right ulna; MHNL 20-149227, fragment of shaπ of right ulna; MHNL 20-149228, fragment of shaπ of leπ humerus + fragment of shaπ of ulna; MHNL 20-149229, incomplete proximal part of leπ humerus; MHNL 20-149231, partial leπ tarsometatarsus; MHNL 20-149234, fragment of shaπ of leπ humerus; MHNL 20-149235, incomplete distal part of leπ ulna; MHNL 20-149236, fragments of shaπ of humerus + fragment of distal end of leπ ulna; MHNL 20-149237, incomplete distal part of leπ ulna; MHNL 20-149238, complete right tibiotarsus; D1-0027B, incomplete cranium; D1-0027C, incomplete cranium with portion of proximal maxilla; D1-0027D, incomplete cranium; D1-0027E, distal end of maxilla; MHNMKhg 10, leπ humerus embedded in matrix. TABLE 1: Comparative measurements of the cranium in Dasornis toliapica and Dasornis emuinus . Lengths in mm. * = estimated measurements; ** = strong dorsoventral flattening.

D. toliapica	D. emuinus
Length (rostral end-prominentia cerebellaris)	59 (BMNH 44096) 63* (OCP.DEK/GE 1043) 65* (OCP.DEK/GE 1238) 66.1 (OCP.DEK/GE 1287) 69 (OCP.DEK/GE 1076) 80* (OCP.DEK/GE 1044) 82* (BMNH A1)	102* (OCP.DEK/GE 1288) 105–110* (based on OCP.DEK/GE 324 and D1-0027A) 110 (BMNH 31929)
Width at rostral end (zona flexoria craniofacialis)	19 (BMNH 44096) 23 (OCP.DEK/GE 1043) 25* (OCP.DEK/GE 1238) 25 (OCP.DEK/GE 1287) 27* (OCP.DEK/GE 1076) 28* (OCP.DEK/GE 1044) 30 (BMNH A1)	36* (OCP.DEK/GE 324) 41* (BMNH 31929)
Minimal width in interorbital region	12 (BMNH 44096) 14 (OCP.DEK/GE 1076, 1238) 15* (OCP.DEK/GE 1287) 16 (OCP.DEK/GE 1044) 18 (BMNH A1)	25.7 (OCP.DEK/GE 1004) 26.8 (OCP.DEK/GE 324) 29* (D1-0027A) 30* (OCP.DEK/GE 1288) 32.1 (BMNH 31929)
Width at processus postorbitales	49* (OCP.DEK/GE 1043) 51* (OCP.DEK/GE 1076) 58* (OCP.DEK/GE 1230) 59.5* (OCP.DEK/GE 1044)	90* (OCP.DEK/GE 1004) 91* (OCP.DEK/GE 324) 95* (D1-0027A) 99* (BMNH A3691) 102* (BMNH 31929)
Lamina parasphenoidalis: width at tuberculi basilares	15.9 (BMNH 44096) 17 (OCP.DEK/GE 1043) 18 (OCP.DEK/GE 1042) 19 (OCP.DEK/GE 1076) 21.5 (OCP.DEK/GE 1044)	35.4 (BMNH A3691) 38* (OCP.DEK/GE 324) 39* (OCP.DEK/GE 1004) 41 (BMNH 31929) 41.5 (D1-0027A)
Height (apex-lamina parasphenoi- dalis)	24 (OCP.DEK/GE 1043) 24.5 (OCP.DEK/GE 1076) 31.5 (OCP.DEK/GE 1044) 32 (OCP.DEK/GE 1042) 33 (BMNH 44096) 41.3 (BMNH A1)	43.4** (D1-0027A) 51.3 (OCP.DEK/GE 1004) 58 (BMNH 31929) 64 (BMNH A3691)
Foramen magnum: width	7 (OCP.DEK/GE 1042) 8.2 (BMNH 44096)	11 (D1-0027A) 11.3 (BMNH 31929)
Foramen magnum: height	7.5* (BMNH 44096) 9 (OCP.DEK/GE 1042)	9** (D1-0027A) 13.3 (BMNH 31929)
Condylus occipitalis: width	6 (OCP.DEK/GE 1042, 1043) 6.2 (OCP.DEK/GE 1076) 7 (OCP.DEK/GE 1044)	11 (D1-0027A) 11.7 (OCP.DEK/GE 324) 12* (OCP.DEK/GE 1288) 12.5 (SMNK-PAL 4017; aπer Mayr, 2008) 12.6 (BMNH 31929)
Processus rostropterygoideus: length	10 (OCP.DEK/GE 1042) 10.5 (OCP.DEK/GE 1044) 12.2 (BMNH 44096)	16.5 (D1-0027A) 18.5 (OCP.DEK/GE 1004)

D. toliapica	D. emuinus
Laterosphenoid hollow: width	12 (OCP.DEK/GE 1043) 13 (OCP.DEK/GE 1042) 14 (OCP.DEK/GE 1076, D1-0027B) 17 (OCP.DEK/GE 1044)	28* (OCP.DEK/GE 324, 1004) 29* (BMNH A3691) 34 (D1-0027A)
Height of maxilla at apertura nasi ossea	20* (OCP.DEK/GE 1185)	—
Maximum width of maxilla at apertura nasi ossea	17* (OCP.DEK/GE 1185)	—
Apertura nasi ossea: length/height	7*/2 (OCP.DEK/GE 1185)	—
Estimated whole length of maxilla	170* (based on BMNH 44096, OCP. DEK/GE 1185 and D1-0027E)	—
Ratio length of maxilla/length of cranium	2/5* (based on estimated length of maxilla and OCP.DEK/GE 1076)	—
Maximum height of mandibula as preserved	17.5 (OCP.DEK/GE 1166)	—

LOCALITIES: Ouled Abdoun Basin ( Morocco ): Sidi Daoui (including Krupp, PH 2, RP13, TS and TZ5 areas); Recette IV; southern Meraa El Arech (Lahou area); Sidi Chennane (M1S2, M8, M11 and R6–R7 areas). STRATIGRAPHIC DISTRIBUTION: Bed IIa, Thanetian, Upper Paleocene; intercalary beds II/I, basal Ypresian, lowermost Eocene; bed I, Ypresian, Lower Eocene. AMENDED DIAGNOSIS (from Harrison and Walker, 1976 ): Albatross-sized odontopterygiform with a wingspan of 2 to 3 m ; differs from Dasornis emuinus in the following features: much smaller size; cranium, sutura frontoparietalis less pronounced, forming barely distinguishable groove; lamina parasphenoidalis caudorostrally longer; caudal extension of cotyla quadratica otici shorter and not twisted ventrally; foramen pneumaticum dorsale larger; laterosphenoid hollow square in shape, bounded medially by conspicuous prominentia lobi optici; tibiotarsus with shorter sulcus musculi fibularis. MEASUREMENTS: See tables 1 and 2 .