1262:
118:
379:. Like in modern flamingos, the region before the nares leading up to the tip is lined with foramina, however uniquely the bone itself is mostly solid and not reticulated like in extant taxa. This trait is consistent across both the juvenile bill illustrated by Olson and Feduccia and the bill of an older individual described by Cheneval. The maxillary keel, located on the underside of the upper bill, is more similar to
389:. Both ML StG 203bis as well as previously recovered bill material show that the keel forms two ridges which eventually broaden and flatten until forming a single ridge that extends to the tip of the bill. The overall shape of the bill is subject to change as the animal grows, as shown by the upper bill material. The juvenile remains show a much shallower curvature, while in adults the bill curves more abruptly.
1492:
294:. Although the type material was only poorly described by Gervais and later lost, other fossils of skull material would eventually be found, specifically the rostral halves of both upper and lower bill. Harrison and Walker, who described one of said bill fragments in 1976, considered the fossils to be distinct enough to warrant a new genus,
656:
held its head more vertically while filtering, as opposed to the upside-down position assumed by modern flamingos. Although the articulation of the lower bill was also found to differ, with the reduction of certain muscles impacting the way the head could be held during feeding and display, it is not
646:
was already a filter feeding animal. In modern taxa its the ventral surface of the upper bill as well as the margins of the lower bill that hold the keratinous lamellae that allow flamingos to filter water in search of prey, while the deep lower bill also holds the enlarged tongue of these birds. The
664:
as well as other fossil birds recovered at Saint-Gérand-le-Puy in an attempt to reconstruct the environment of the area. With over half of the fossil birds analysed by
Cheneval preferring brackish water and no exclusively freshwater species present, he concludes that the area was likely covered by
392:
The lower bill shows a strong dorsal curvature matching the way that the upper bill curves downward with ridges on the sides typical for flamingos. As expected for members of its family, the lower bill is much deeper than the upper and strongly mediolaterally compressed. When put together with the
367:
the nares stand out due to their proximity to the point where the bill begins its downward curvature, extending beyond the beginning of the curvature and almost approaching the inflection point. The frontal bones also resemble modern flamingos, with high ridges just behind the nares and shallow
651:
is much shallower than in modern species, as well as the shallower, less curved lower bill, supports the idea that the filtering mechanism in this genus was not yet as well developed as in the flamingos of today. Specifically, the curvature of the bill may have meant that
437:
was already developed by the Late
Oligocene and saw very little change until today, with the other modern flamingo genera simply showing an even greater specialisation and increased filtering efficiency towards small organisms like
665:
brackish water. Later research found that Saint-Gérand-le-Puy was a complex of lacustrine environments undergoing repeated change between wet and dry conditions, causing brackish conditions through evaporation, as indicated by
312:
in 1978. Another upper bill was figured by Olson and
Feduccia and assigned to a juvenile individual in 1980, but not described in detail. Eventually, another In 2015, another skull (specimen ML StG 203bis) was described from
397:
bill was much less curved overall than in any modern flamingo, which matches the narrower maxillary keel. In size the upper bills FSL 442292 and NMB MA 9594 both indicate a size similar to the modern
868:
Cheneval, J. (1989). "Fossil bird study, and paleoecological and paleoenvironmental consequences: Example from the Saint-Gérand-le-Puy deposits (lower miocene, Allier, France)".
413:
lies outside the crown-group of flamingos, which diverged during the
Pliocene to Pleistocene. Although previous publications have argued that the similarities between
317:. This skull consisted of most of the cranium and the adjacent caudal region of the bill, making it the first cranial material since the lost type material.
1556:
921:
704:
Harrison, C. J. O.; Walker, C. A. (1976). "Cranial material of
Oligocene and Miocene flamingos: with a description of new species from Africa".
1584:
1261:
251:. Although generally similar, it subtly differs in the curvature of the bill and the size of the ventral keel of the maxilla, both signs that
247:. Despite being one of the oldest known members of the flamingo family, it already shows a skull remarkably similar to that of the modern
1604:
1599:
290:. Due to this superficial similarity, it was assumed that the fossils belonged to an extinct species of said genus, then named
764:
914:
117:
1589:
1594:
907:
284:
on the basis of a nearly complete skull which showed a similar morphology to modern flamingos of the genus
1511:
899:
314:
429:, Torres and colleagues argue that the similarity is due to the plesiomorphic state of the bill seen in
731:
Olson, S.L.; Feduccia, A. (1980). "Relationships and evolution of flamingos (Aves: Phoenicopteridae)".
1543:
1198:
1122:
1074:
877:
822:
811:"A multi-locus inference of the evolutionary diversification of extant flamingos (Phoenicopteridae)"
1491:
1291:
1279:
1233:
169:
769:(Aves, Phoenicopteridae) informs the evolution of the highly specialized filter feeding apparatus"
791:
240:
112:
34:
296:
1329:
1019:
850:
1398:
1206:
1130:
1081:
1031:
885:
840:
830:
783:
713:
398:
248:
1472:
376:
421:
indicate that the two were more closely related to one another than to the deeper billed
881:
826:
1462:
1448:
1441:
1387:
993:
981:
845:
810:
677:
577:
560:
335:
286:
1578:
1455:
889:
341:
71:
795:
1534:
1355:
1173:
1164:
511:
281:
260:
205:
809:
Torres, C.R.; Ogawa, L.M.; Gillingham, M.A.; Ferrari, B.; van Tuinen, M. (2014).
1431:
1318:
1138:
1106:
1009:
464:
385:
383:, if even shallower than seen in either the modern species or the extinct genus
46:
1377:
1367:
1340:
1307:
1183:
1115:
1067:
787:
683:
359:
are elongated, but more similar to those of modern flamingos than to those of
347:
91:
56:
1047:
835:
305:
129:
96:
40:
854:
1528:
1217:
1191:
1148:
1091:
1057:
954:
935:
667:
236:
232:
179:
149:
86:
81:
66:
61:
51:
30:
1561:
1410:
966:
439:
333:
species with their large and compact bills, setting it apart from both
101:
76:
763:
Torres, C. R.; De Pietri, V. L.; Louchart, A.; Van Tuinen, M. (2015).
948:
717:
487:
255:
was not yet as adapted towards filter feeding as modern species are.
244:
139:
1505:
931:
352:
1548:
960:
642:
The close resemblance to modern flamingo species indicates that
159:
1509:
1277:
979:
903:
765:"New cranial material of the earliest filter feeding flamingo
706:
Bulletin of the
British Museum (Natural History), Geology
393:
upper bill, it becomes noticeable that the silhouette of
351:, a phoenicopteriform with a straight, pointed beak. The
657:
yet clear how great the effect would have been exactly.
345:
with their smaller, bulbous bills and the contemporary
1468:
1437:
1427:
1417:
1406:
1383:
1373:
1363:
1352:
1336:
1325:
1314:
1303:
1213:
1179:
1144:
1087:
1063:
1053:
1043:
1015:
1005:
675:
shared this environment with palaeolodids including
1518:
1397:
1351:
1290:
1232:
1163:
1105:
1030:
992:
375:, is broad and shallow, not spoon-shaped as in the
363:, in which the nares take up much of the bill. In
329:broadly agrees with the morphology seen in modern
259:inhabited brackish lakes alongside the more basal
870:Palaeogeography, Palaeoclimatology, Palaeoecology
433:. This shows that the general feeding ecology of
915:
280:was first reported in 1852 by paleontologist
8:
1506:
1296:
1287:
1274:
1160:
1102:
1036:
998:
989:
976:
922:
908:
900:
20:
844:
834:
758:
756:
754:
752:
750:
748:
746:
409:The anatomy of the skull indicates that
696:
300:. Unbenownst to them however, the name
304:was already occupied by a species of
7:
733:Smithsonian Contributions to Zoology
776:Organisms Diversity & Evolution
552:
527:
503:
480:
455:
445:
371:The upper bill, much like that of
14:
263:. It contains the single species
1490:
1260:
116:
368:fossae just above the orbits.
308:. Kashin renamed the genus to
1:
1585:Fossil taxa described in 1852
325:The known fossil material of
890:10.1016/0031-0182(89)90010-2
938:and their extinct relatives
269:, first described in 1852.
1621:
1605:Oligocene birds of Europe
1488:
1299:
1286:
1273:
1258:
1039:
1001:
988:
975:
943:
788:10.1007/s13127-015-0209-7
574:
557:
550:
532:
525:
508:
501:
485:
478:
460:
453:
211:
204:
113:Scientific classification
111:
23:
1600:Prehistoric bird genera
836:10.1186/1471-2148-14-36
292:Phoenicopterus croizeti
243:periods in what is now
231:is an extinct genus of
214:Phoenicopterus croizeti
647:fact that the keel of
235:that lived during the
767:Harrisonavis croizeti
266:Harrisonavis croizeti
499:Phoenicopteriformes
273:Discovery and naming
1292:Phoenicopteriformes
1280:Phoenicopteriformes
1234:Phoenicopteriformes
882:1989PPP....73..295C
827:2014BMCEE..14...36T
315:Saint-Gérand-le-Puy
170:Phoenicopteriformes
1590:Fossils of France
1572:
1571:
1512:Taxon identifiers
1503:
1502:
1499:
1498:
1486:
1485:
1482:
1481:
1269:
1268:
1256:
1255:
1252:
1251:
1228:
1227:
1159:
1158:
1101:
1100:
634:
633:
625:
624:
616:
615:
607:
606:
598:
597:
589:
588:
523:Phoenicopteridae
224:
223:
200:
1612:
1595:Phoenicopteridae
1565:
1564:
1552:
1551:
1539:
1538:
1537:
1507:
1494:
1470:
1439:
1429:
1419:
1408:
1399:Phoenicopteridae
1385:
1375:
1365:
1354:
1338:
1327:
1316:
1305:
1297:
1288:
1275:
1264:
1215:
1181:
1161:
1146:
1103:
1089:
1065:
1055:
1045:
1037:
1032:Podicipediformes
1017:
1007:
999:
990:
977:
924:
917:
910:
901:
894:
893:
876:(3–4): 295–309.
865:
859:
858:
848:
838:
806:
800:
799:
773:
760:
741:
740:
728:
722:
721:
718:10.5962/p.313816
701:
553:
528:
504:
488:Podicipediformes
481:
456:
446:
399:greater flamingo
249:greater flamingo
198:
191:
180:Phoenicopteridae
121:
120:
106:
43:
29:Temporal range:
21:
1620:
1619:
1615:
1614:
1613:
1611:
1610:
1609:
1575:
1574:
1573:
1568:
1560:
1555:
1547:
1542:
1533:
1532:
1527:
1514:
1504:
1495:
1478:
1473:Xenorhynchopsis
1393:
1347:
1282:
1265:
1248:
1224:
1155:
1097:
1026:
984:
971:
939:
928:
898:
897:
867:
866:
862:
808:
807:
803:
771:
762:
761:
744:
730:
729:
725:
703:
702:
698:
693:
640:
635:
626:
617:
608:
599:
590:
407:
377:lesser flamingo
323:
275:
220:
217:
197:
189:
115:
107:
105:
104:
99:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
38:
37:
27:
17:
12:
11:
5:
1618:
1616:
1608:
1607:
1602:
1597:
1592:
1587:
1577:
1576:
1570:
1569:
1567:
1566:
1553:
1540:
1524:
1522:
1516:
1515:
1510:
1501:
1500:
1497:
1496:
1489:
1487:
1484:
1483:
1480:
1479:
1477:
1476:
1466:
1463:Phoenicoparrus
1459:
1452:
1449:Phoenicopterus
1445:
1442:Phoeniconotius
1435:
1425:
1415:
1403:
1401:
1395:
1394:
1392:
1391:
1388:Megapaloelodus
1381:
1371:
1360:
1358:
1349:
1348:
1346:
1345:
1334:
1323:
1312:
1300:
1294:
1284:
1283:
1278:
1271:
1270:
1267:
1266:
1259:
1257:
1254:
1253:
1250:
1249:
1247:
1246:
1245:
1244:
1238:
1236:
1230:
1229:
1226:
1225:
1223:
1222:
1211:
1203:
1195:
1188:
1177:
1169:
1167:
1157:
1156:
1154:
1153:
1142:
1135:
1127:
1119:
1111:
1109:
1099:
1098:
1096:
1095:
1085:
1078:
1071:
1061:
1051:
1040:
1034:
1028:
1027:
1025:
1024:
1013:
1002:
996:
994:Mirandornithes
986:
985:
982:Mirandornithes
980:
973:
972:
970:
969:
963:
957:
951:
944:
941:
940:
929:
927:
926:
919:
912:
904:
896:
895:
860:
801:
782:(3): 609–618.
742:
723:
712:(4): 305–314.
695:
694:
692:
689:
678:Megapaloelodus
660:Cheneval used
639:
636:
632:
631:
628:
627:
623:
622:
619:
618:
614:
613:
610:
609:
605:
604:
601:
600:
596:
595:
592:
591:
587:
586:
583:
582:
578:Phoenicoparrus
573:
570:
569:
566:
565:
561:Phoenicopterus
556:
551:
549:
546:
545:
542:
541:
531:
526:
524:
520:
519:
516:
515:
507:
502:
500:
496:
495:
492:
491:
484:
479:
477:
474:
473:
470:
469:
459:
454:
452:
444:
435:Phoenicopterus
431:Phoenicopterus
423:Phoenicoparrus
419:Phoenicopterus
406:
403:
381:Phoenicopterus
373:Phoenicopterus
336:Phoenicoparrus
331:Phoenicopterus
322:
319:
287:Phoenicopterus
274:
271:
237:Late Oligocene
222:
221:
218:
209:
208:
202:
201:
187:
183:
182:
177:
173:
172:
167:
163:
162:
157:
153:
152:
147:
143:
142:
137:
133:
132:
127:
123:
122:
109:
108:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
44:
28:
16:Genus of birds
15:
13:
10:
9:
6:
4:
3:
2:
1617:
1606:
1603:
1601:
1598:
1596:
1593:
1591:
1588:
1586:
1583:
1582:
1580:
1563:
1558:
1554:
1550:
1545:
1541:
1536:
1530:
1526:
1525:
1523:
1521:
1517:
1513:
1508:
1493:
1475:
1474:
1467:
1465:
1464:
1460:
1458:
1457:
1456:Phoeniconaias
1453:
1451:
1450:
1446:
1444:
1443:
1436:
1434:
1433:
1426:
1424:
1423:
1416:
1413:
1412:
1405:
1404:
1402:
1400:
1396:
1390:
1389:
1382:
1380:
1379:
1372:
1370:
1369:
1362:
1361:
1359:
1357:
1350:
1343:
1342:
1335:
1332:
1331:
1324:
1321:
1320:
1313:
1310:
1309:
1302:
1301:
1298:
1295:
1293:
1289:
1285:
1281:
1276:
1272:
1263:
1242:
1241:
1240:
1239:
1237:
1235:
1231:
1220:
1219:
1212:
1209:
1208:
1204:
1201:
1200:
1199:Poliocephalus
1196:
1194:
1193:
1189:
1186:
1185:
1178:
1176:
1175:
1171:
1170:
1168:
1166:
1162:
1151:
1150:
1143:
1141:
1140:
1136:
1133:
1132:
1128:
1125:
1124:
1123:Poliocephalus
1120:
1118:
1117:
1113:
1112:
1110:
1108:
1104:
1094:
1093:
1086:
1084:
1083:
1079:
1077:
1076:
1075:Poliocephalus
1072:
1070:
1069:
1062:
1060:
1059:
1052:
1050:
1049:
1042:
1041:
1038:
1035:
1033:
1029:
1022:
1021:
1014:
1012:
1011:
1004:
1003:
1000:
997:
995:
991:
987:
983:
978:
974:
968:
964:
962:
958:
956:
952:
950:
946:
945:
942:
937:
933:
925:
920:
918:
913:
911:
906:
905:
902:
891:
887:
883:
879:
875:
871:
864:
861:
856:
852:
847:
842:
837:
832:
828:
824:
820:
816:
815:BMC Evol Biol
812:
805:
802:
797:
793:
789:
785:
781:
777:
770:
768:
759:
757:
755:
753:
751:
749:
747:
743:
738:
734:
727:
724:
719:
715:
711:
707:
700:
697:
690:
688:
686:
685:
680:
679:
674:
670:
669:
663:
658:
655:
650:
645:
637:
630:
629:
621:
620:
612:
611:
603:
602:
594:
593:
585:
584:
581:
580:
579:
572:
571:
568:
567:
564:
563:
562:
555:
554:
548:
547:
544:
543:
540:
539:
538:
530:
529:
522:
521:
518:
517:
514:
513:
506:
505:
498:
497:
494:
493:
490:
489:
483:
482:
476:
475:
472:
471:
468:
467:
466:
458:
457:
451:
450:Mirandornites
448:
447:
443:
441:
436:
432:
428:
427:Phoeniconaias
424:
420:
416:
412:
404:
402:
400:
396:
395:Harrisonavis'
390:
388:
387:
382:
378:
374:
369:
366:
362:
358:
354:
350:
349:
344:
343:
342:Phoeniconaias
338:
337:
332:
328:
320:
318:
316:
311:
307:
303:
299:
298:
293:
289:
288:
283:
279:
272:
270:
268:
267:
262:
258:
254:
250:
246:
242:
241:Early Miocene
238:
234:
230:
229:
219:Gervais, 1852
216:
215:
210:
207:
203:
196:
195:
188:
185:
184:
181:
178:
175:
174:
171:
168:
165:
164:
161:
158:
155:
154:
151:
148:
145:
144:
141:
138:
135:
134:
131:
128:
125:
124:
119:
114:
110:
103:
98:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
42:
36:
32:
26:
22:
19:
1520:Harrisonavis
1519:
1471:
1461:
1454:
1447:
1440:
1430:
1422:Harrisonavis
1421:
1420:
1409:
1386:
1376:
1366:
1356:Palaelodidae
1339:
1328:
1317:
1306:
1216:
1205:
1197:
1190:
1182:
1174:Aechmophorus
1172:
1165:Podicipedini
1147:
1137:
1129:
1121:
1114:
1090:
1080:
1073:
1066:
1056:
1046:
1018:
1008:
873:
869:
863:
818:
814:
804:
779:
775:
766:
736:
732:
726:
709:
705:
699:
682:
676:
673:Harrisonavis
672:
671:gastropods.
666:
662:Harrisonavis
661:
659:
654:Harrisonavis
653:
649:Harrisonavis
648:
644:Harrisonavis
643:
641:
576:
575:
559:
558:
537:Harrisonavis
536:
535:
533:
512:Palaelodidae
509:
486:
463:
461:
449:
434:
430:
426:
422:
418:
415:Harrisonavis
414:
411:Harrisonavis
410:
408:
394:
391:
384:
380:
372:
370:
365:Harrisonavis
364:
360:
357:Harrisonavis
356:
346:
340:
334:
330:
327:Harrisonavis
326:
324:
310:Harrisonavis
309:
301:
295:
291:
285:
282:Paul Gervais
278:Harrisonavis
277:
276:
265:
264:
261:Palaelodidae
257:Harrisonavis
256:
253:Harrisonavis
252:
228:Harrisonavis
227:
226:
225:
213:
212:
206:Type species
199:Kashin, 1978
194:Harrisonavis
193:
192:
25:Harrisonavis
24:
18:
1432:Leakeyornis
1319:Juncitarsus
1243:See below ↓
1139:Tachybaptus
1107:Podilymbini
1010:Juncitarsus
465:Juncitarsus
386:Leakeyornis
321:Description
39:28–13
1579:Categories
1535:Q110269247
1378:Palaelodus
1368:Adelalopus
1341:Scaniornis
1308:Agnopterus
1184:Pliolymbus
1116:Podilymbus
1068:Pliolymbus
930:Genera of
691:References
684:Palaelodus
361:Palaelodus
348:Palaelodus
35:Aquitanian
1207:Rollandia
1131:Rollandia
1082:Rollandia
1048:Miobaptus
947:Kingdom:
936:flamingos
405:Phylogeny
306:millipede
302:Gervaisia
297:Gervaisia
136:Kingdom:
130:Eukaryota
1549:60973493
1529:Wikidata
1330:Kashinia
1218:Thiornis
1192:Podiceps
1149:Thiornis
1092:Thiornis
1058:Miodytes
1020:Kashinia
955:Chordata
953:Phylum:
949:Animalia
855:24580860
796:18198929
668:Hydrobia
233:flamingo
176:Family:
150:Chordata
146:Phylum:
140:Animalia
126:Domain:
31:Chattian
1562:4850510
1411:Elornis
967:Neoaves
965:Clade:
959:Class:
878:Bibcode
846:4016592
823:Bibcode
739:: 1–84.
638:Ecology
440:diatoms
186:Genus:
166:Order:
156:Class:
932:grebes
853:
843:
821:(36).
794:
245:France
792:S2CID
772:(PDF)
353:nares
1557:GBIF
961:Aves
851:PMID
681:and
425:and
417:and
339:and
239:and
160:Aves
47:PreꞒ
1544:EoL
886:doi
841:PMC
831:doi
784:doi
737:316
714:doi
355:of
1581::
1559::
1546::
1531::
934:,
884:.
874:73
872:.
849:.
839:.
829:.
819:14
817:.
813:.
790:.
780:15
778:.
774:.
745:^
735:.
710:27
708:.
687:.
510:†
462:†
442:.
401:.
97:Pg
41:Ma
1469:†
1438:†
1428:†
1418:†
1414:?
1407:†
1384:†
1374:†
1364:†
1353:†
1344:?
1337:†
1333:?
1326:†
1322:?
1315:†
1311:?
1304:†
1221:?
1214:†
1210:?
1202:?
1187:?
1180:†
1152:?
1145:†
1134:?
1126:?
1088:†
1064:†
1054:†
1044:†
1023:?
1016:†
1006:†
923:e
916:t
909:v
892:.
888::
880::
857:.
833::
825::
798:.
786::
720:.
716::
534:†
190:†
102:N
92:K
87:J
82:T
77:P
72:C
67:D
62:S
57:O
52:Ꞓ
33:-
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