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tree. However, the homology between the ornithischian filaments and the theropods' proto-feathers is not obvious. If the homology is supported, the consequence is that the common ancestor of both saurischians and ornithischians were covered by feather-like structures, and that groups for which skin
755:
The filamentous integumentary structures are preserved on three areas of the fossil: in one patch just below the neck, another one on the back, and the largest one above the tail. The hollow filaments are parallel to each other and are singular with no evidence of branching. They also appear to be
844:
to dinosaurs. The results supported the hypothesis that scales are plesiomorphic to dinosaurs. While it is true that feather beta keratin is present in crocodilian scales in embryonic development, it fails to support the maximum-likelihood of protofeathers being plesiomorphic.
519:
is classified as a heterodontosaurid, a group of small ornithischian dinosaur characterized by a slender body, long tail and a pair of enlarged canine-like tusks. They were herbivorous or possibly omnivorous. Until the discovery of
786:. The estimated length of the integumentary structures on the tail is about 60 mm which is seven times the height of a caudal vertebra. Their length and hollow nature argue against of them being subdermal structures such as
451:, as well as remains of long, singular and unbranched filamentous integumentary structures. The holotype is from a subadult individual that probably measured 70 cm in length based on the proportions of the related
803:
were only secondarily featherless. If the homology is not supported, it would indicate that these filamentous dermal structures evolved independently in saurischians and ornithischians, as well as in other
1333:
917:
Liu Y.-Q. Kuang H.-W., Jiang X.-J., Peng N., Xu H. & Sun H.-Y. (2012). "Timing of the earliest known feathered dinosaurs and transitional pterosaurs older than the Jehol Biota."
836:, the filaments were round in cross section, and therefore closer in structure to the earliest forms of feathers predicted by developmental models. A study published in the journal
2406:
1038:
865:
Zheng, Xiao-Ting; You, Hai-Lu; Xu, Xing; Dong, Zhi-Ming (19 March 2009). "An Early
Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures".
2504:
1155:
1256:"Molecular evolution and expression of archosaurian β-keratins: diversification and expansion of archosaurian β-keratins and the origin of feather β-keratins"
1052:
Mayr, Gerald; Peters, D. Stephan; Plodowski, Gerhard; Vogel, Olaf (August 2002). "Bristle-like integumentary structures at the tail of the horned dinosaur
465:
had unusual proportions compared to other heterodontosaurids. The head was large and the legs and tail were long, but the neck and forelimbs were short.
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1370:
812:. The authors (in supplementary information to their primary article) noted that discovery of similar filamentous structures in the theropod
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fibers. Such dermal structures have previously been reported only in derived theropods and ornithischians, and their discovery in
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structures on the back, tail and neck of the specimen. The similarity of these structures with those found on some
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457:
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1554:
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and raises the possibility that the earliest dinosaurs and their ancestors were covered with homologous
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Another specimen, IVPP V17090, was described in 2012. At least four other specimens remain undescribed.
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from the
Quantou Formation (Cretaceous: Aptian–Cenomanian) of Jilin Province, north-eastern China"
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relatively rigid, making them more analogous to the integumentary structures found on the tail of
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995:"The postcranial osteology and phylogenetic position of the small ornithischian dinosaur
936:"Taxonomy, morphology, masticatory function and phylogeny of heterodontosaurid dinosaurs"
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365:. The fossil was collected at a locality transliterated as Linglengta or Linglongta. Lu
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filamentous structures that can be considered primitive feathers ("proto-feathers").
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1151:"A new feather type in a nonavian theropod and the early evolution of feathers"
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STMN 26-3 consists of an incomplete skeleton preserving a partial skull and
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Barrett, Paul M.; Evans, David C.; Campione, Nicolás E. (2015-06-30).
361:, STMN 26-3 was initially reported as being from the Early Cretaceous
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Specimen IVPP V17090, muzzle, hand, feet and tail framed in red
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are most similar to the singular unbranched proto-feathers of
1338:- a fuzzy dinosaur that makes the origin of feathers fuzzier"
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extends the existence of such structures further down in the
369:, 2010, reported that these beds were actually part of the
1149:
Xu, Xing; Zheng Xiao-ting; You, Hai-lu (20 January 2009).
1254:
Greenwold, Matthew J.; Sawyer, Roger H. (June 6, 2013).
840:
rigorously tested the hypothesis that protofeathers are
2407:
List of stratigraphic units with ornithischian tracks
820:
are homologous with feathers. Both the filaments of
524:, known members of the group were restricted to the
2438:
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919:Palaeogeography, Palaeoclimatology, Palaeoecology
1156:Proceedings of the National Academy of Sciences
538:of the US, and possibly one additional genus (
1364:
8:
1207:"Evolution of dinosaur epidermal structures"
1037:: CS1 maint: multiple names: authors list (
832:, however, the filaments were flattened. In
1066:(8). Heidelberg: Springer Berlin: 361–365.
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2085:
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1749:
1530:
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1371:
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816:bolstered the idea that the structures on
116:
20:
1230:
1178:
1168:
1022:
989:Richard J. Butler, Jin Liyong, Chen Jun,
965:
947:
770:. Among the theropods, the structures in
377:period at least 158.5 million years ago.
828:were long, singular, and unbranched. In
766:structures found in avian and non-avian
544:) from the Early Cretaceous of England.
1109:Currie, Philip J.; Pei-ji Chen (2001).
854:
1030:
551:below follows the analysis by Butler
7:
1200:
1198:
929:
927:
677:
614:
607:
566:
558:
1123:Canadian Journal of Earth Sciences
1115:from Liaoning, northeastern China"
14:
1260:Journal of Experimental Zoology B
799:impression are known such as the
528:of South Africa, with one genus (
415:vertebrae, nearly complete right
333:whose remains were discovered in
2178:
2171:
2164:
2158:
2151:
1313:
1024:10.1111/j.1475-4983.2011.01046.x
392:The holotype skeleton, STMN 26-3
306:Shandong Tianyu Museum of Nature
134:
2574:Late Jurassic dinosaurs of Asia
471:has a row of long, filamentous
423:, the proximal end of the left
1:
2589:Fossil taxa described in 2009
1130:(12). NRC Canada: 1705–1727.
921:(advance online publication).
934:Sereno, Paul (2012-03-10).
2640:
2599:Taxa named by Dong Zhiming
562:Heterodontosauridae
16:Extinct genus of dinosaurs
2614:Monotypic dinosaur genera
2202:
2191:
2148:
1853:
1752:
1533:
1529:
1514:
1501:
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1386:
1080:10.1007/s00114-002-0339-6
699:
682:
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632:
619:
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588:
571:
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315:is housed) is an extinct
263:
256:
131:Scientific classification
129:
124:
115:
23:
2579:Paleontology in Liaoning
949:10.3897/zookeys.226.2840
2604:Jurassic ornithischians
1170:10.1073/pnas.0810055106
1136:10.1139/cjes-38-12-1705
458:Heterodontosaurus tucki
373:, dating from the late
2609:Ornithischians of Asia
1223:10.1098/rsbl.2015.0229
997:Changchunsaurus parvus
752:
513:
393:
329:. The only species is
2594:Taxa named by Xu Xing
2531:Paleobiology Database
1113:Sinosauropteryx prima
824:and the filaments of
750:
502:
391:
270:Tianyulong confuciusi
2564:Ornithischian genera
371:Tiaojishan Formation
2584:Feathered dinosaurs
1676:Heterodontosauridae
1280:10.1002/jez.b.22514
1272:2013JEZB..320..393G
1072:2002NW.....89..361M
1059:Naturwissenschaften
1015:2011Palgy..54..667B
889:10.1038/nature07856
881:2009Natur.458..333Z
210:Heterodontosauridae
2624:Dinosaurs of China
2559:Heterodontosaurids
2386:Hypsilophodontidae
2124:Pachycephalosauria
2113:Pachycephalosauria
1332:(March 18, 2009).
762:than to the proto-
753:
514:
503:Skull diagrams of
394:
249:T. confuciusi
2546:
2545:
2518:Open Tree of Life
2432:Taxon identifiers
2423:
2422:
2419:
2418:
2415:
2414:
2354:Serendipaceratops
2187:
2186:
2146:
2145:
2142:
2141:
2138:
2137:
2134:
2133:
1838:
1837:
1834:
1833:
1706:Heterodontosaurus
1497:
1496:
1493:
1492:
1489:
1488:
875:(7236): 333–336.
739:
738:
730:
729:
721:
720:
712:
711:
664:
663:
636:Heterodontosaurus
506:Heterodontosaurus
339:Liaoning Province
321:heterodontosaurid
285:
284:
237:
2631:
2569:Oxfordian genera
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2538:
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2513:
2512:
2500:
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1933:Thescelosauridae
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1503:
1449:
1440:
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1388:
1373:
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1359:
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1345:
1344:on May 15, 2012.
1340:. Archived from
1323:
1321:Dinosaurs portal
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991:Pascal Godefroit
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335:Jianchang County
304:; named for the
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29:Temporal range:
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2376:Phytodinosauria
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2198:
2183:
2170:
2163:
2150:
2130:
2107:
2081:Marginocephalia
2075:
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1949:Changchunsaurus
1927:
1847:
1830:
1807:
1784:
1739:
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1523:
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1459:Avemetatarsalia
1444:Avemetatarsalia
1434:
1419:Avemetatarsalia
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1392:
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1328:
1319:
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1217:(6): 20150229.
1211:Biology Letters
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838:Biology Letters
777:Sinosauropteryx
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479:suggests their
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2619:Jurassic China
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2400:Other articles
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2391:Jeholosauridae
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2371:Ornithoscelida
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2322:Incertae sedis
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2032:Thescelosaurus
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2014:
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1942:Albertadromeus
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1880:Kulindadromeus
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1845:Neornithischia
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1772:Scutellosaurus
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1307:External links
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1301:
1266:(6): 393–405.
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1101:
1054:Psittacosaurus
1044:
1009:(3): 667–683.
981:
942:(226): 1–225.
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759:Psittacosaurus
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2480:
2476:
2471:
2465:
2461:
2456:
2450:
2446:
2445:
2443:
2441:
2437:
2433:
2428:
2408:
2405:
2404:
2402:
2398:
2392:
2389:
2387:
2384:
2382:
2379:
2377:
2374:
2372:
2369:
2368:
2366:
2362:
2356:
2355:
2351:
2349:
2348:
2344:
2341:
2340:
2339:Dravidosaurus
2336:
2334:
2333:
2329:
2328:
2326:
2324:
2323:
2318:
2312:
2311:
2307:
2305:
2304:
2300:
2298:
2297:
2293:
2291:
2290:
2289:Syngonosaurus
2286:
2284:
2283:
2279:
2277:
2276:
2272:
2270:
2269:
2265:
2263:
2262:
2258:
2256:
2255:
2251:
2249:
2248:
2244:
2242:
2241:
2237:
2235:
2234:
2230:
2228:
2227:
2223:
2221:
2220:
2216:
2215:
2213:
2211:
2210:
2205:
2201:
2194:
2190:
2181:
2177:
2174:
2167:
2161:
2154:
2125:
2121:
2120:
2119:
2118:
2116:
2114:
2110:
2102:
2098:
2097:
2096:
2095:
2093:
2091:
2087:
2084:
2082:
2078:
2070:
2066:
2065:
2064:
2063:
2061:
2059:
2055:
2049:
2048:
2047:Zephyrosaurus
2044:
2041:
2040:
2036:
2034:
2033:
2029:
2027:
2026:
2022:
2020:
2019:
2018:Oryctodromeus
2015:
2013:
2012:
2008:
2006:
2005:
2004:Nevadadromeus
2001:
1998:
1997:
1993:
1990:
1989:
1988:Koreanosaurus
1985:
1982:
1981:
1977:
1974:
1973:
1969:
1967:
1966:
1962:
1959:
1958:
1954:
1951:
1950:
1946:
1944:
1943:
1939:
1938:
1936:
1934:
1930:
1924:
1923:
1919:
1917:
1916:
1912:
1910:
1909:
1905:
1903:
1902:
1898:
1896:
1895:
1891:
1889:
1888:
1884:
1882:
1881:
1877:
1875:
1874:
1873:Hypsilophodon
1870:
1868:
1867:
1866:Hexinlusaurus
1863:
1861:
1860:
1856:
1855:
1852:
1849:
1846:
1841:
1825:
1821:
1820:
1819:
1818:
1816:
1814:
1810:
1802:
1798:
1797:
1796:
1795:
1793:
1791:
1787:
1781:
1780:
1776:
1774:
1773:
1769:
1767:
1766:
1765:Scelidosaurus
1762:
1760:
1759:
1755:
1754:
1751:
1748:
1746:
1742:
1736:
1735:
1731:
1729:
1728:
1724:
1722:
1721:
1717:
1715:
1714:
1710:
1708:
1707:
1703:
1701:
1700:
1696:
1694:
1693:
1689:
1687:
1686:
1685:Abrictosaurus
1682:
1681:
1679:
1677:
1673:
1666:
1665:
1661:
1658:
1657:
1653:
1650:
1649:
1645:
1642:
1641:
1637:
1634:
1633:
1629:
1626:
1625:
1621:
1618:
1617:
1613:
1611:
1610:
1609:Lesothosaurus
1606:
1604:
1603:
1602:Laquintasaura
1599:
1596:
1595:
1591:
1588:
1587:
1583:
1580:
1579:
1575:
1572:
1571:
1570:Eucoelophysis
1567:
1565:
1564:
1560:
1557:
1556:
1552:
1549:
1548:
1544:
1541:
1540:
1536:
1535:
1532:
1528:
1525:
1522:
1517:
1513:
1509:
1504:
1500:
1479:
1478:
1477:
1476:
1474:
1472:
1468:
1460:
1456:
1455:
1454:
1453:
1450:
1447:
1445:
1441:
1437:
1429:
1425:
1422:
1420:
1416:
1413:
1411:
1407:
1405:
1401:
1399:
1395:
1394:
1389:
1385:
1381:
1374:
1369:
1367:
1362:
1360:
1355:
1354:
1351:
1343:
1339:
1337:
1331:
1327:
1326:
1322:
1311:
1306:
1297:
1293:
1289:
1285:
1281:
1277:
1273:
1269:
1265:
1261:
1257:
1250:
1247:
1242:
1238:
1233:
1228:
1224:
1220:
1216:
1212:
1208:
1201:
1199:
1195:
1190:
1186:
1181:
1176:
1171:
1166:
1162:
1158:
1157:
1152:
1145:
1142:
1137:
1133:
1129:
1125:
1124:
1116:
1114:
1105:
1102:
1097:
1093:
1089:
1085:
1081:
1077:
1073:
1069:
1065:
1061:
1060:
1055:
1048:
1045:
1040:
1034:
1025:
1020:
1016:
1012:
1008:
1004:
1003:Palaeontology
1000:
998:
992:
985:
982:
977:
973:
968:
963:
959:
955:
950:
945:
941:
937:
930:
928:
924:
920:
914:
911:
906:
902:
898:
894:
890:
886:
882:
878:
874:
870:
869:
861:
859:
855:
848:
846:
843:
842:plesiomorphic
839:
835:
831:
830:Beipiaosaurus
827:
826:Beipiaosaurus
823:
819:
815:
814:Beipiaosaurus
811:
807:
802:
797:
793:
789:
785:
784:
783:Beipiaosaurus
779:
778:
773:
769:
765:
761:
760:
749:
742:
735:
734:
726:
725:
717:
716:
708:
707:
704:
703:
697:
696:
693:
692:
689:
688:
687:
680:
679:
673:
672:
669:
668:
660:
659:
656:
655:
654:
647:
646:
643:
642:
639:
638:
637:
630:
629:
626:
625:
622:
617:
616:
610:
609:
603:
602:
599:
598:
595:
594:
593:
592:Abrictosaurus
586:
585:
582:
581:
578:
577:
576:
569:
568:
561:
560:
556:
554:
550:
545:
543:
542:
537:
536:Late Jurassic
533:
532:
527:
523:
518:
512:
508:
507:
501:
494:
492:
490:
486:
482:
478:
474:
473:integumentary
470:
466:
464:
460:
459:
454:
453:South African
450:
446:
442:
438:
434:
430:
426:
422:
418:
414:
410:
406:
403:
399:
390:
383:
381:
378:
376:
372:
368:
364:
360:
356:
348:
346:
344:
340:
336:
332:
328:
325:
324:ornithischian
322:
318:
314:
311:
307:
303:
299:
295:
291:
290:
279:
273:
271:
262:
259:
258:Binomial name
255:
251:
250:
244:
241:
240:
236:
232:
226:
225:
218:
215:
214:
211:
205:
202:
201:
198:
192:
189:
186:
185:
182:
179:
176:
173:
172:
169:
166:
163:
162:
159:
156:
153:
152:
149:
146:
143:
142:
137:
132:
128:
123:
119:
114:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
39:
32:
31:Late Jurassic
26:
22:
19:
2439:
2381:Silesauridae
2352:
2345:
2337:
2332:Ajkaceratops
2330:
2320:
2308:
2301:
2294:
2287:
2280:
2275:Notoceratops
2273:
2266:
2261:Gongbusaurus
2259:
2254:Geranosaurus
2252:
2245:
2238:
2231:
2224:
2217:
2209:Nomina dubia
2207:
2157:
2045:
2037:
2030:
2025:Parksosaurus
2023:
2016:
2009:
2002:
1994:
1986:
1980:Jeholosaurus
1978:
1970:
1963:
1955:
1947:
1940:
1920:
1913:
1908:Sanxiasaurus
1906:
1899:
1892:
1887:Minimocursor
1885:
1878:
1871:
1864:
1857:
1801:Ankylosauria
1790:Ankylosauria
1777:
1770:
1763:
1756:
1733:
1732:
1725:
1718:
1711:
1704:
1697:
1690:
1683:
1664:Technosaurus
1662:
1656:Soumyasaurus
1654:
1646:
1638:
1632:Pisanosaurus
1630:
1622:
1614:
1607:
1600:
1592:
1586:Ignotosaurus
1584:
1576:
1568:
1561:
1553:
1545:
1537:
1521:Ornithischia
1508:Ornithischia
1471:Ornithischia
1423:
1414:
1380:Ornithischia
1342:the original
1335:
1263:
1259:
1249:
1214:
1210:
1163:(3): 832–4.
1160:
1154:
1144:
1127:
1121:
1112:
1111:"Anatomy of
1104:
1063:
1057:
1053:
1047:
1033:cite journal
1006:
1002:
996:
984:
939:
918:
913:
872:
866:
837:
833:
829:
825:
821:
817:
813:
808:such as the
796:phylogenetic
791:
781:
775:
771:
757:
754:
743:Paleobiology
701:
700:
684:
683:
651:
650:
634:
633:
621:NHM RU A100
620:
590:
589:
573:
572:
552:
546:
539:
529:
521:
516:
515:
510:
504:
468:
467:
462:
456:
439:, the right
395:
379:
366:
358:
352:
330:
301:
288:
287:
286:
277:
269:
264:
248:
247:
230:
223:
222:
197:Ornithischia
187:
174:
24:
18:
2464:Wikispecies
2310:Trimucrodon
2282:Siluosaurus
2240:Fabrosaurus
2233:Craspedodon
2226:Bienosaurus
2069:Ornithopoda
2058:Ornithopoda
1957:Changmiania
1922:Yandusaurus
1859:Agilisaurus
1824:Stegosauria
1813:Stegosauria
1745:Thyreophora
1624:Lutungutali
1594:Kwanasaurus
1547:Asilisaurus
1539:Amanasaurus
751:Restoration
534:) from the
461:. However,
384:Description
363:Jehol group
36:158.5
2553:Categories
2470:Tianyulong
2440:Tianyulong
2364:Other taxa
2303:Tichosteus
2296:Tatisaurus
2101:Ceratopsia
2090:Ceratopsia
2039:Yueosaurus
2011:Orodromeus
1996:Nanosaurus
1915:Xiaosaurus
1894:Nanosaurus
1779:Yuxisaurus
1758:Emausaurus
1734:Tianyulong
1727:Pegomastax
1713:Lycorhinus
1699:Fruitadens
1648:Silesaurus
1640:Sacisaurus
1616:Lewisuchus
1480:see below↓
1428:Dinosauria
1410:Sauropsida
1336:Tianyulong
849:References
834:Tianyulong
822:Tianyulong
818:Tianyulong
810:pterosaurs
806:archosaurs
792:Tianyulong
772:Tianyulong
702:Tianyulong
686:Fruitadens
653:Lycorhinus
531:Fruitadens
522:Tianyulong
517:Tianyulong
511:Tianyulong
469:Tianyulong
463:Tianyulong
427:, partial
400:, partial
359:Tianyulong
337:, Western
308:where the
302:tiānyǔlóng
289:Tianyulong
224:Tianyulong
181:Dinosauria
25:Tianyulong
2268:Laosaurus
1901:Phyllodon
1692:Echinodon
1578:Gamatavus
1396:Kingdom:
1288:1552-5015
958:1313-2970
801:sauropods
768:theropods
575:Echinodon
549:cladogram
541:Echinodon
477:theropods
405:vertebrae
402:presacral
242:Species:
154:Kingdom:
148:Eukaryota
2484:10657572
2449:Wikidata
2219:Alocodon
2197:See also
1720:Manidens
1563:Eocursor
1555:Diodorus
1404:Chordata
1402:Phylum:
1398:Animalia
1296:23744807
1241:26041865
1189:19139401
1096:17781405
1088:12435037
993:(2011).
976:23166462
897:19295609
788:collagen
555:, 2011:
485:feathers
481:homology
455:species
411:–middle
409:proximal
398:mandible
375:Jurassic
355:holotype
327:dinosaur
310:holotype
203:Family:
168:Chordata
164:Phylum:
158:Animalia
144:Domain:
2523:4126676
2510:1478649
2497:4965667
2455:Q739132
2347:Jakapil
1408:Class:
1330:Ed Yong
1268:Bibcode
1232:4528472
1180:2630069
1068:Bibcode
1011:Bibcode
967:3491919
940:ZooKeys
905:4423110
877:Bibcode
764:feather
435:, both
431:, both
419:, both
417:scapula
349:History
296:: 天宇龍;
294:Chinese
267:†
246:†
216:Genus:
105:↓
2536:143495
1294:
1286:
1239:
1229:
1187:
1177:
1094:
1086:
974:
964:
956:
903:
895:
868:Nature
553:et al.
489:dermal
445:fibula
437:femora
433:ischia
421:humeri
413:caudal
367:et al.
313:fossil
298:Pinyin
280:, 2009
278:et al.
276:Zheng
231:et al.
229:Zheng
2505:IRMNG
1424:Clade
1415:Clade
1118:(PDF)
1092:S2CID
901:S2CID
483:with
441:tibia
429:pubes
343:China
317:genus
188:Clade
175:Clade
2492:GBIF
2122:see
2099:see
2067:see
1972:Haya
1965:Fona
1822:see
1799:see
1457:see
1292:PMID
1284:ISSN
1237:PMID
1185:PMID
1084:PMID
1039:link
972:PMID
954:ISSN
893:PMID
780:and
547:The
509:and
447:and
443:and
425:ulna
353:The
235:2009
44:PreꞒ
2479:EoL
1276:doi
1264:320
1227:PMC
1219:doi
1175:PMC
1165:doi
1161:106
1132:doi
1076:doi
1056:".
1019:doi
962:PMC
944:doi
885:doi
873:458
449:pes
357:of
319:of
2555::
2533::
2520::
2507::
2494::
2481::
2466::
2451::
1426::
1417::
1290:.
1282:.
1274:.
1262:.
1258:.
1235:.
1225:.
1215:11
1213:.
1209:.
1197:^
1183:.
1173:.
1159:.
1153:.
1128:38
1126:.
1120:.
1090:.
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1074:.
1064:89
1062:.
1035:}}
1031:{{
1017:.
1007:54
1005:.
1001:.
970:.
960:.
952:.
938:.
926:^
899:.
891:.
883:.
871:.
857:^
407:,
345:.
341:,
300::
233:,
190::
177::
94:Pg
38:Ma
33:,
2342:?
2042:?
1999:?
1991:?
1983:?
1975:?
1960:?
1952:?
1667:?
1659:?
1651:?
1643:?
1635:?
1627:?
1619:?
1597:?
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1372:e
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1278::
1270::
1243:.
1221::
1191:.
1167::
1138:.
1134::
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1078::
1070::
1041:)
1027:.
1021::
1013::
978:.
946::
907:.
887::
879::
292:(
220:†
207:†
194:†
99:N
89:K
84:J
79:T
74:P
69:C
64:D
59:S
54:O
49:Ꞓ
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