2082:
129:
610:
353:
649:
104:
636:
restudied the holotype and described new features in 1922. He described many of these features from a fragment of the snout and lower jaw. Although this fossil was well preserved, it was not found in the same block of marl as other parts of the specimen. This fossil was reexamined in 2003 and was
522:
457:
to have both a pineal and a parapineal eye, as the only other vertebrates that have both are the jawless lampreys. In most vertebrates, the pineal gland forms the parietal eye, however, in lepidosaurs, it is formed from the parapineal gland. This implies that
370:
had a long, pointed snout and nostrils placed farther back in the skull than most lizards and a tail that was almost twice as long as the body. Although similar in appearance to
555:
was not a species of lizard, noting that the humerus was "undoubtedly" nonreptilian. Leidy even pointed out similarities between the bone and those of birds in 1873.
536:
Although his first description was brief, Leidy studied the genus thoroughly and provided illustrations in an 1873 paper. In this paper, Leidy called it
1994:"The anatomy, phylogenetic relationships, and autecology of the carnivorous lizard "Saniwa" feisti Stritzke, 1983 from the Eocene of Messel, Germany"
1844:
2159:
1380:
773:
was described from the Green River
Formation of Wyoming in 2007. It preserves soft tissues like scales, cartilage between bones and in the
510:
to Leidy because it was "used by one of the Indian tribes of the Upper
Missouri for a rock-lizard." Leidy saw a close similarity between
1330:
942:
570:
in 1872 for several species of extinct lizards in the western United States. He never published a full description of these lizards, and
1832:. Report of the United States Geological Survey of the Territories. Vol. 1. United States Government Printing Office. p. 358.
2213:
1921:
Augé, Marc Louis; Annelise FOLIE, undefined; Smith, Richard; PhéLizon, Alain; Paul GIGASE†, undefined; Smith, Thierry (2022-08-02).
2146:
2243:
2238:
2233:
398:
1640:
1562:
1704:
1590:
1612:
116:
1899:
1923:"Revision of the oldest varanid, Saniwa orsmaelensis Dollo, 1923, from the earliest Eocene of northwest Europe"
1641:"Re-assessment of varanid evolution based on new data from Saniwa ensidens Leidy, 1870 (Squamata, Reptilia)"
1993:
1922:
1859:
1691:. Handbook of Sensory Physiology. Vol. 7 / 3 / 3 B. Springer, Berlin, Heidelberg. pp. 113–140.
417:
and the other from the parapineal gland. The parietal eye is a light-sensitive structure present in the
471:
1648:
2223:
1554:
128:
2218:
1886:
1599:
2228:
2190:
2099:
1720:
Smith, Krister T.; Bhullar, Bhart-Anjan S.; Köhler, Gunther; Habersetzer, Jörg (2 April 2018).
2164:
2185:
2177:
1687:
Dodt, Eberhard (1973). "The
Parietal Eye (Pineal and Parietal Organs) of Lower Vertebrates".
563:
313:
272:
1735:
632:, including the underside of the skull and parts of the vertebrae. American paleontologist
214:
1968:
n. sp., ein
Varanide (Lacertilia, Reptilia) aus dem Mittel-Eozän von Messel bei Darmstadt"
8:
2087:
674:
2039:
Philosophical
Transactions of the Royal Society of London. Series B, Biological Sciences
1739:
628:
was prepared by removing marl from around the bones. This revealed many new features of
2059:
2034:
2015:
1903:
1807:
1669:
1616:
778:
721:. Unlike the Argentine fossils, the Belgian remains represent a definite occurrence of
633:
259:
123:
1722:"The Only Known Jawed Vertebrate with Four Eyes and the Bauplan of the Pineal Complex"
2172:
2064:
2019:
1944:
1907:
1761:
1753:
1700:
1620:
1558:
551:
and some isolated dorsal vertebrae. In 1918, Baron G. J. de Fejérváry suggested that
503:
309:
1811:
1673:
2137:
2054:
2046:
2005:
1963:
1934:
1895:
1799:
1743:
1692:
1661:
1653:
1608:
479:
446:
1803:
494:
was the first extinct lizard to be named from North
America. The first remains of
2010:
1939:
1783:
1726:
1548:
756:
751:
654:
402:
1696:
2122:
2035:"A new stem-varanid lizard (Reptilia, Squamata) from the early Eocene of China"
1457:
793:
has been recognized as a close relative of living monitor lizards in the genus
575:
340:
1748:
1721:
366:
measured 1.3 to 2.1 m (4.3 to 6.9 ft). Like other varanid lizards,
2207:
1948:
1757:
1398:
1109:
997:
454:
391:
386:
bone beneath the eye that extended farther forward, and a suture between the
328:
61:
2068:
2050:
1765:
810:
802:
690:
638:
515:
483:
430:
414:
410:
387:
321:
237:
210:
609:
2131:
1312:
1287:
1255:
1059:
1039:
1015:
962:
908:
706:
338:, is recognised from remains found in Europe. It is a close relative of
36:
1830:
Contributions to the extinct vertebrate fauna of the
Western Territories
445:. It plays an important role in geographical orientation and regulating
2151:
1845:"Contributions to a monography on the fossil Varanidae and Megalanidae"
840:
738:
426:
371:
352:
81:
46:
1665:
1232:
1091:
985:
818:
294:
190:
140:
86:
2093:
2033:
Dong L, Wang YQ, Zhao Q, Vasilyan D, Wang Y, Evans SE (March 2022).
2116:
857:
614:
180:
160:
76:
71:
56:
51:
41:
1657:
705:
in 1923, but because its naming was informal, it was designated a
673:, although all are fragmentary. In 1899, Argentine paleontologist
1474:
1153:
774:
702:
686:
669:
Fossils from many other parts of the world have been assigned to
548:
442:
438:
418:
317:
170:
91:
66:
648:
1920:
1613:
10.1666/pleo0022-3360(2007)081[0643:TAOTFV]2.0.CO;2
521:
379:
305:
301:
297:
150:
103:
1719:
693:, because the material cannot be assigned with confidence to
434:
383:
290:
327:
was described in 1870 as the first fossil lizard known from
1900:
10.1666/0022-3360(2003)077<0393:HSEOSE>2.0.CO;2
1638:
1595:
Leidy, 1870, based on a newly discovered complete skeleton"
499:
422:
590:, but all other species have remained distinct, including
1876:
Caldwell, M.W. (March 2003). "Holotype snout elements of
1547:
Pianka, Eric R.; King, Dennis; King, Ruth Allen (2004).
733:
with remains of the species also reported from France. "
1852:
Annales
Historico-Naturales Musei Nationalis Hungarici
825:(2008) that shows a sister-group relationship between
1639:
Conrad, J.; Rieppel, O.; Grande, L. (December 2008).
781:. The individual is thought to have been a juvenile.
2077:
2032:
1132:
1105:
1055:
1011:
958:
938:
904:
879:
854:
486:. Later that year, Leidy described the type species
244:
229:
378:had many primitive traits, including teeth on its
1588:
713:was later suggested to be either synonymous with
2205:
1991:
813:analyses, meaning it is more closely related to
624:In the 1920s, much of the holotype specimen of
1788:Leidy, an extinct varanid lizard from Wyoming"
1546:
1992:Smith, Krister T.; Habersetzer, Jörg (2021).
745:is no longer considered to be a species of
689:rocks in Argentina. It is now considered a
344:, the genus that includes monitor lizards.
1591:"The anatomy of the fossil varanid lizard
797:. It is a member of the family Varanidae.
725:outside North America. A 2022 study found
102:
2058:
2009:
1938:
1842:
1747:
717:or a different, indeterminate species of
1961:
1875:
647:
608:
525:Leidy's illustrations of the humerus of
520:
351:
1792:Proceedings of the U.S. National Museum
1781:
1777:
1775:
1634:
1632:
1630:
769:A complete and articulated skeleton of
397:A study in 2018 by scientists from the
304:epoch. It is known from well-preserved
2206:
1823:
1821:
1584:
1582:
1580:
1578:
1576:
1574:
749:, but is placed in the separate genus
729:to be a distinct and valid species of
465:
394:that was straight rather than curved.
2098:
2097:
1827:
1589:Rieppel, O.; Grande, L. (July 2007).
759:, which is more distantly related to
1955:
1869:
1836:
1772:
1686:
1627:
613:The vertebrae and limb bones of the
1818:
1571:
817:than any other varanid. Below is a
13:
2026:
540:. He also named a second species,
482:, and gave them to paleontologist
14:
2255:
784:
498:were preserved as black bones in
2214:Eocene reptiles of North America
2080:
127:
34:
1985:
1914:
490:on the basis of these fossils.
117:Field Museum of Natural History
1713:
1680:
1540:
413:, one that developed from the
399:Senckenberg Research Institute
347:
1:
2244:Fossil taxa described in 1870
1804:10.5479/si.00963801.60-2418.1
1550:Varanoid Lizards of the World
1534:
789:Since its first description,
2011:10.5852/cr-palevol2021v20a23
1940:10.5852/cr-palevol2022v21a25
1884:sp. indet. (Xenosauridae)".
506:. Hayden suggested the name
470:In 1870, American geologist
7:
1843:Fejérváry, G.J. de (1918).
1697:10.1007/978-3-642-65495-4_4
1689:Visual Centers in the Brain
1399:Proplatynotia longirostrata
547:, on the basis of a broken
474:found the first fossils of
10:
2260:
2239:Taxa named by Joseph Leidy
2234:Prehistoric reptile genera
737:was named from the Eocene
562:, American paleontologist
472:Ferdinand Vandeveer Hayden
462:reevolved the pineal eye.
2106:
1858:: 341–467. Archived from
1749:10.1016/j.cub.2018.02.021
1649:American Museum Novitates
1471:
1453:
1446:
1428:
1421:
1394:
1376:
1369:
1362:
1326:
1308:
1301:
1283:
1276:
1269:
1251:
1244:
1150:
1130:
1123:
1103:
1096:
1053:
1036:
1029:
1009:
1002:
990:
956:
936:
929:
922:
902:
895:
877:
870:
852:
845:
677:named another species of
265:
258:
226:
221:
124:Scientific classification
122:
110:
101:
23:
1555:Indiana University Press
1381:Paravaranus angustifrons
16:Extinct genus of lizards
1975:Senckenbergiana Lethaea
1887:Journal of Paleontology
1600:Journal of Paleontology
1235:from Dong et al. 2022.
801:is often placed as the
574:was later considered a
558:Soon after Leidy named
2051:10.1098/rstb.2021.0041
1998:Comptes Rendus Palevol
1927:Comptes Rendus Palevol
1784:"A new description of
1782:Gilmore, C.W. (1922).
1331:Saniwides mongoliensis
666:
621:
533:
360:
314:Green River Formations
300:that lived during the
2186:Paleobiology Database
1962:Stritzke, R. (1983).
1557:. pp. 33 to 38.
741:in Germany in 1983. "
651:
637:found to belong to a
612:
586:was synonymized with
564:Othniel Charles Marsh
529:and the vertebrae of
524:
502:that was part of the
355:
320:, United States. The
1288:Telmasaurus grangeri
449:and annual rhythms.
331:. A second species,
2088:Paleontology portal
1740:2018CBio...28E1101S
1734:(7): 1101–1107.e2.
701:was described from
675:Florentino Ameghino
466:History and species
2045:(1847): 20210041.
1880:reassigned to cf.
1828:Leidy, J. (1873).
1458:Archaeovaranus lii
1313:Aiolosaurus oriens
755:within the family
667:
634:Charles W. Gilmore
622:
566:erected the genus
534:
453:is the only known
361:
2201:
2200:
2173:Open Tree of Life
2100:Taxon identifiers
1865:on 26 March 2012.
1531:
1530:
1522:
1521:
1513:
1512:
1504:
1503:
1495:
1494:
1486:
1485:
1410:
1409:
1351:
1350:
1342:
1341:
1228:
1227:
1219:
1218:
1210:
1209:
1201:
1200:
1192:
1191:
1183:
1182:
1174:
1173:
1165:
1164:
1080:
1079:
1071:
1070:
974:
973:
504:Bridger Formation
478:near the town of
374:monitor lizards,
282:
281:
276:
252:
241:
217:
2251:
2194:
2193:
2181:
2180:
2168:
2167:
2155:
2154:
2142:
2141:
2140:
2127:
2126:
2125:
2095:
2094:
2090:
2085:
2084:
2083:
2073:
2072:
2062:
2030:
2024:
2023:
2013:
1989:
1983:
1982:
1972:
1959:
1953:
1952:
1942:
1918:
1912:
1911:
1873:
1867:
1866:
1864:
1849:
1840:
1834:
1833:
1825:
1816:
1815:
1779:
1770:
1769:
1751:
1717:
1711:
1710:
1684:
1678:
1677:
1645:
1636:
1625:
1624:
1586:
1569:
1568:
1544:
1449:
1448:
1424:
1423:
1372:
1371:
1365:
1364:
1304:
1303:
1279:
1278:
1272:
1271:
1247:
1246:
1239:
1238:
1134:
1126:
1125:
1107:
1099:
1098:
1057:
1032:
1031:
1013:
1005:
1004:
993:
992:
960:
940:
932:
931:
925:
924:
906:
898:
897:
881:
873:
872:
856:
848:
847:
837:
836:
480:Granger, Wyoming
455:jawed vertebrate
421:, most lizards,
271:
250:
246:
235:
231:
209:
202:
132:
131:
106:
96:
33:
29:Temporal range:
21:
20:
2259:
2258:
2254:
2253:
2252:
2250:
2249:
2248:
2224:Monitor lizards
2204:
2203:
2202:
2197:
2189:
2184:
2176:
2171:
2163:
2158:
2150:
2145:
2136:
2135:
2130:
2121:
2120:
2115:
2102:
2086:
2081:
2079:
2076:
2031:
2027:
2004:(23): 441–506.
1990:
1986:
1981:(5–6): 497–508.
1970:
1960:
1956:
1933:(25): 511–529.
1919:
1915:
1878:Saniwa ensidens
1874:
1870:
1862:
1847:
1841:
1837:
1826:
1819:
1786:Saniwa ensidens
1780:
1773:
1727:Current Biology
1718:
1714:
1707:
1685:
1681:
1643:
1637:
1628:
1593:Saniwa ensidens
1587:
1572:
1565:
1545:
1541:
1537:
1532:
1523:
1514:
1505:
1496:
1487:
1433:Saniwa ensidens
1411:
1352:
1343:
1229:
1220:
1211:
1202:
1193:
1184:
1175:
1166:
1137:Saniwa ensidens
1081:
1072:
975:
883:"Saniwa" feisti
827:Saniwa ensidens
799:Saniwa ensidens
787:
777:, and even the
757:Palaeovaranidae
752:Paranecrosaurus
727:S. orsmaelensis
711:S. orsmaelensis
699:S. orsmaelensis
655:Paranecrosaurus
514:and the living
488:Saniwa ensidens
468:
403:Yale University
350:
248:S. orsmaelensis
208:
200:
126:
113:Saniwa ensidens
97:
95:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
39:
31:
30:
27:
17:
12:
11:
5:
2257:
2247:
2246:
2241:
2236:
2231:
2226:
2221:
2219:Eocene lizards
2216:
2199:
2198:
2196:
2195:
2182:
2169:
2156:
2143:
2128:
2112:
2110:
2104:
2103:
2092:
2091:
2075:
2074:
2025:
1984:
1954:
1913:
1894:(2): 393–396.
1868:
1835:
1817:
1771:
1712:
1705:
1679:
1652:(3630): 1–15.
1626:
1607:(4): 643–665.
1570:
1564:978-0253343666
1563:
1538:
1536:
1533:
1529:
1528:
1525:
1524:
1520:
1519:
1516:
1515:
1511:
1510:
1507:
1506:
1502:
1501:
1498:
1497:
1493:
1492:
1489:
1488:
1484:
1483:
1480:
1479:
1470:
1467:
1466:
1463:
1462:
1452:
1447:
1445:
1442:
1441:
1438:
1437:
1427:
1422:
1420:
1417:
1416:
1413:
1412:
1408:
1407:
1404:
1403:
1393:
1390:
1389:
1386:
1385:
1375:
1370:
1368:
1363:
1361:
1358:
1357:
1354:
1353:
1349:
1348:
1345:
1344:
1340:
1339:
1336:
1335:
1325:
1322:
1321:
1318:
1317:
1307:
1302:
1300:
1297:
1296:
1293:
1292:
1282:
1277:
1275:
1270:
1268:
1265:
1264:
1261:
1260:
1250:
1245:
1243:
1237:
1226:
1225:
1222:
1221:
1217:
1216:
1213:
1212:
1208:
1207:
1204:
1203:
1199:
1198:
1195:
1194:
1190:
1189:
1186:
1185:
1181:
1180:
1177:
1176:
1172:
1171:
1168:
1167:
1163:
1162:
1159:
1158:
1149:
1146:
1145:
1142:
1141:
1129:
1124:
1122:
1119:
1118:
1115:
1114:
1102:
1097:
1095:
1087:
1086:
1083:
1082:
1078:
1077:
1074:
1073:
1069:
1068:
1065:
1064:
1052:
1049:
1048:
1045:
1044:
1035:
1030:
1028:
1025:
1024:
1021:
1020:
1008:
1003:
1001:
991:
989:
981:
980:
977:
976:
972:
971:
968:
967:
955:
952:
951:
948:
947:
935:
930:
928:
923:
921:
918:
917:
914:
913:
901:
896:
894:
891:
890:
887:
886:
876:
871:
869:
866:
865:
862:
861:
851:
846:
844:
835:
786:
785:Classification
783:
582:. The species
576:junior synonym
467:
464:
392:parietal bones
349:
346:
289:is an extinct
280:
279:
278:
277:
263:
262:
256:
255:
254:
253:
242:
224:
223:
219:
218:
198:
194:
193:
188:
184:
183:
178:
174:
173:
168:
164:
163:
158:
154:
153:
148:
144:
143:
138:
134:
133:
120:
119:
108:
107:
99:
98:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
40:
35:
28:
15:
9:
6:
4:
3:
2:
2256:
2245:
2242:
2240:
2237:
2235:
2232:
2230:
2229:Lizard genera
2227:
2225:
2222:
2220:
2217:
2215:
2212:
2211:
2209:
2192:
2187:
2183:
2179:
2174:
2170:
2166:
2161:
2157:
2153:
2148:
2144:
2139:
2133:
2129:
2124:
2118:
2114:
2113:
2111:
2109:
2105:
2101:
2096:
2089:
2078:
2070:
2066:
2061:
2056:
2052:
2048:
2044:
2040:
2036:
2029:
2021:
2017:
2012:
2007:
2003:
2000:(in French).
1999:
1995:
1988:
1980:
1976:
1969:
1967:
1966:Saniwa feisti
1958:
1950:
1946:
1941:
1936:
1932:
1928:
1924:
1917:
1909:
1905:
1901:
1897:
1893:
1889:
1888:
1883:
1879:
1872:
1861:
1857:
1853:
1846:
1839:
1831:
1824:
1822:
1813:
1809:
1805:
1801:
1797:
1793:
1789:
1787:
1778:
1776:
1767:
1763:
1759:
1755:
1750:
1745:
1741:
1737:
1733:
1729:
1728:
1723:
1716:
1708:
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1019:
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998:Lanthanotinae
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983:
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411:parietal eyes
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356:The skull of
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26:
22:
19:
2107:
2042:
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2001:
1997:
1987:
1978:
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1965:
1957:
1930:
1926:
1916:
1891:
1885:
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1877:
1871:
1860:the original
1855:
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1838:
1829:
1798:(23): 1–28.
1795:
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961:
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941:
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882:
878:
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830:
826:
822:
821:from Conrad
814:
811:phylogenetic
806:
803:sister taxon
798:
794:
790:
788:
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764:
760:
750:
746:
742:
734:
730:
726:
722:
718:
714:
710:
698:
694:
691:dubious name
683:S. australis
682:
678:
670:
668:
663:
659:
658:, formerly "
653:
652:A fossil of
642:
641:lizard, not
629:
625:
623:
618:
617:specimen of
604:T. paucidens
603:
599:
595:
591:
587:
583:
579:
571:
567:
559:
557:
552:
544:
541:
537:
535:
530:
526:
516:Nile monitor
511:
507:
495:
491:
487:
484:Joseph Leidy
475:
469:
459:
450:
443:jawless fish
415:pineal gland
406:
396:
375:
367:
363:
362:
357:
339:
336:orsmaelensis
335:
332:
324:
322:type species
285:
284:
283:
268:
247:
232:
204:
203:
112:
111:Skeleton of
24:
18:
2132:Wikispecies
1256:Ovoo gurvel
1231:Below is a
1060:Cherminotus
1040:Lanthanotus
1016:Aiolosaurus
963:Telmasaurus
909:Necrosaurus
771:S. ensidens
715:S. ensidens
626:S. ensidens
619:S. ensidens
588:S. ensidens
584:T. leptodus
572:Thinosaurus
568:Thinosaurus
531:S. ensidens
496:S. ensidens
427:salamanders
358:S. ensidens
348:Description
325:S. ensidens
269:Thinosaurus
251:Dollo, 1923
233:S. ensidens
2208:Categories
1535:References
1242:Varanidae
841:Varanoidea
743:S." feisti
739:Messel Pit
735:S." feisti
707:naked name
639:xenosaurid
600:T. grandis
429:, certain
2020:236259655
1949:1777-571X
1908:129541157
1758:0960-9822
1666:2246/5939
1621:130800851
1233:cladogram
1092:Varaninae
986:Varanidae
943:Saniwides
858:Mosasaurs
819:cladogram
596:T. crassa
592:T. agilis
447:circadian
431:bony fish
191:Varanidae
147:Kingdom:
141:Eukaryota
2123:Q3948438
2117:Wikidata
2069:35125002
1812:22291809
1766:29614279
1674:83550662
615:holotype
553:S. major
527:S. major
439:lampreys
409:had two
260:Synonyms
222:Species
187:Family:
181:Squamata
171:Reptilia
161:Chordata
157:Phylum:
151:Animalia
137:Domain:
2178:4119731
2165:1295712
2152:4820733
2060:8819366
1736:Bibcode
1475:Varanus
1154:Varanus
1094:
1000:
988:
843:
831:Varanus
815:Varanus
807:Varanus
795:Varanus
779:trachea
775:sternum
765:Saniwa.
761:Varanus
731:Saniwa,
703:Belgium
687:Miocene
549:humerus
419:tuatara
388:frontal
341:Varanus
318:Wyoming
310:Bridger
306:fossils
295:varanid
197:Genus:
177:Order:
167:Class:
115:in the
2138:Saniwa
2108:Saniwa
2067:
2057:
2018:
1947:
1906:
1882:Restes
1810:
1764:
1756:
1703:
1672:
1619:
1561:
1090:
996:
984:
823:et al.
791:Saniwa
747:Saniwa
723:Saniwa
719:Saniwa
695:Saniwa
679:Saniwa
671:Saniwa
664:feisti
660:Saniwa
643:Saniwa
630:Saniwa
602:, and
580:Saniwa
560:Saniwa
542:Saniwa
538:Saniwa
512:Saniwa
508:Saniwa
492:Saniwa
476:Saniwa
460:Saniwa
451:Saniwa
435:sharks
407:Saniwa
405:found
380:palate
376:Saniwa
372:extant
368:Saniwa
364:Saniwa
302:Eocene
298:lizard
286:Saniwa
275:, 1872
205:Saniwa
32:Eocene
25:Saniwa
2191:38041
2160:IRMNG
2016:S2CID
1971:(PDF)
1904:S2CID
1863:(PDF)
1848:(PDF)
1808:S2CID
1670:S2CID
1644:(PDF)
1617:S2CID
763:than
545:major
423:frogs
384:jugal
291:genus
273:Marsh
211:Leidy
2147:GBIF
2065:PMID
1945:ISSN
1762:PMID
1754:ISSN
1701:ISBN
1559:ISBN
1110:Ovoo
829:and
500:marl
437:and
401:and
390:and
382:, a
312:and
238:type
215:1870
37:Preęž’
2055:PMC
2047:doi
2043:377
2006:doi
1935:doi
1896:doi
1800:doi
1744:doi
1693:doi
1662:hdl
1654:doi
1609:doi
809:in
805:of
578:of
316:of
293:of
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2014:.
2002:20
1996:.
1979:64
1977:.
1973:.
1943:.
1931:21
1929:.
1925:.
1902:.
1892:77
1890:.
1856:16
1854:.
1850:.
1820:^
1806:.
1796:60
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1774:^
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1732:28
1730:.
1724:.
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1629:^
1615:.
1605:81
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1573:^
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833::
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697:.
681:,
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645:.
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598:,
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425:,
333:S.
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1951:.
1937::
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