1709:. They were distinguished from chimaeras by a pair of large spines that emanate from their chests, the presence of large scales and plates, tooth-like beak plates, and the typical bone-enhanced placoderm eyeball. They were distinguished from other placoderms due to differences in the anatomy of their skulls, and due to patterns on the skull plates and thoracic plates that are unique to this order. From what can be inferred from the mouthplates of fossil specimens, acanthothoracids were shellfish hunters ecologically similar to modern-day chimaeras. Competition with their relatives, the ptyctodont placoderms, may have been one of the main reasons for the acanthothoracids' extinction prior to the mid-Devonian extinction event.
5373:
5403:
1343:
5414:
3960:
1426:
1762:
3064:
1585:
1954:
154:
5398:
4707:
5380:
5408:
1639:
1355:
5392:
911:
1683:
5192:
3078:
1568:("leaf scales") were flattened placoderms found throughout the world. Like other flattened placoderms they were bottom-dwelling predators that ambushed prey. Unlike other flattened placoderms, they were freshwater fish. Their armour was made of whole plates, rather than the numerous tubercles and scales of Petalichthyida. The eyes were on the sides of the head, unlike visual bottom-dwelling predators, such as
927:
124:
1839:
716:
1554:
1494:
5386:
1050:
979:
1936:
951:
1718:
1674:. However, through comparisons of skull anatomies, rhenanids are now considered to be the sister group of the antiarchs. When rhenanids die, their "mosaics" come apart, and it has been suggested that the rarity of rhenanids in the fossil record reflects postmortem disassociation, and is not an actual rarity of the species.
1796:, the pseudopetalichthids' placement within Placodermi is suspect. The matter is not easy to resolve because there are no complete, undamaged and articulated specimens. The anatomical studies done on the crushed specimens that have been found indicate that if they are placoderms, they may be a group more advanced than the
1014:, given a combination of the supposed inherent superiority of bony fish and the presumed sluggishness of placoderms. With more accurate summaries of prehistoric organisms, it is now thought that they systematically died out as marine and freshwater ecologies suffered from the environmental catastrophes of the
888:
hoxd13, the origin of the mating organs in placoderms most likely relied on different sets of hox genes and were structures that developed further down the body as an extra and independent pair of appendages, but which during development turned into body parts used for reproduction only. Because they
1622:, while those of ptyctodontids are made of bone; the craniums of holocephalians are similar to sharks, while those of ptyctodontids are similar to those of other placoderms; and, most importantly, that holocephalians have true teeth, while ptyctodonts have beak-like tooth plates. Ptyctodontids were
1878:
with thin, strap-like pectoral fins. Similar to those of the
Rhenanida, its armour was a complex mosaic of small, scale-like tubercles. The shoulder joints of its armour are similar to other placoderms, and there are superficial similarities in skull plates, and even more superficial similarities
1203:
Paleontologists and placoderm specialists suspect that the scarcity of placoderms in the
Silurian fossil record is due to placoderms' living in environments unconducive to fossil preservation, rather than a genuine scarcity. This hypothesis helps to explain the placoderms' seemingly instantaneous
1456:
was the mouth, and the opening for the anal siphon was on the other side of the body, as opposed to having both oral and anal siphons together at one end. The front portions of their bodies were heavily armoured, to the point of literally resembling a box with eyes, with the sometimes scaled,
1274:. Once the specimens had been completely ground away (and so destroyed), he made enlarged, three-dimensional models of the skulls to examine the anatomical details more thoroughly. Many other placoderm specialists thought that Stensiö was trying to shoehorn placoderms into a relationship with
1665:
with large, upturned mouths that lived in marine environments. The rhenanids were once presumed to be the most primitive, or at least the closest to the ancestral placoderm, as their armour was made of unfused components—a mosaic of tubercles—as opposed to the solidified plates of "advanced"
1088:), Silurian-aged placoderm specimens consisted of fragments. Some of them have been tentatively identified as antiarch or arthrodire due to histological similarities; and many of them have not yet been formally described or even named. The most commonly cited example of a Silurian placoderm,
3681:
1753:. Soon after the petalichthids' diversification, they went into decline. Because they had compressed body forms, it is supposed they were bottom-dwellers that pursued or ambushed smaller fish. Their diet is not clear, as none of the fossil specimens found have preserved mouth parts.
1485:, the limbs were long and had elbow-like joints. The function of the limbs is still not perfectly understood, but most hypothesize that they helped their owners pull themselves across the substrate, as well as allowing their owners to bury themselves into the substrate.
1746:
are among the best known. There was an independent diversification event that occurred in what is now
Southern China, producing a handful of unique genera that were once placed in their own order, "Quasipetalichthyida", named after the first discovered species there,
1094:
of
Silurian China and possibly Vietnam, is known only from a few fragments that currently defy attempts to place them in any of the recognized placoderm orders. So far, only three officially described Silurian placoderms are known from more than scraps:
3706:
672:, were bottom-dwellers. In particular, the antiarchs, with their highly modified, jointed bony pectoral fins, were highly successful inhabitants of Middle-Late Devonian freshwater and shallow marine habitats, with the Middle to Late Devonian
843:), had eggs which were fertilized internally, the mother providing nourishment to the embryo and giving birth to live young. With this discovery, the placoderm became the oldest vertebrate known to have given birth to live young ("
737:, was 3.5–4.1 metres (11–13 ft) long, and is presumed to have had a large distribution, as its remains have been found in Europe, North America and possibly Morocco. Some paleontologists regard it as the world's first
1732:("thin-plated fish") were small, flattened placoderms, typified by their splayed fins and numerous tubercles that decorated all of the plates and scales of their armour. They reached a peak in diversity during the
1613:
or the ancestors of holocephalians. Anatomical examinations of whole fossil specimens have shown that the similarities between these two groups are superficial. The major differences were that holocephalians have
1776:("false petalichthyids") is a group of elongated, possibly flattened fishes comprising three, poorly preserved and poorly studied genera. It is known only from rare fossils in Lower Devonian strata in
1285:
Much later, the exquisitely preserved placoderm fossils from Gogo reef changed the picture again. They showed that placoderms shared anatomical features not only with chondrichthyans but with other
1322:. These orders are considered to be basal or primitive groups within Placodermi, though their precise placement within the class remains unsure. Fossils of both are currently known only from the
1270:. He took fossil specimens with well-preserved skulls and ground them away, one tenth of a millimeter at a time. After each layer had been removed, he made an imprint of the next surface in
1227:
boundary, the Late
Devonian extinctions. The remaining species then died out during the end-Devonian extinction; not a single placoderm species has been confirmed to have survived into the
1289:
groups as well. For example, Gogo placoderms show separate bones for the nasal capsules as in gnathostomes; in both sharks and bony fish those bones are incorporated into the braincase.
1381:. They had a movable joint between armour surrounding the head and body. As the lower jaw moved down, the head shield moved, allowing for a larger opening. All arthrodires, save for
4032:
Zhu, Min; Ahlberg, Per E.; Pan, Zhaohui; Zhu, Youan; Qiao, Tuo; Zhao, Wenjin; Jia, Liantao; Lu, Jing (21 October 2016). "A Silurian maxillate placoderm illuminates jaw evolution".
3696:
3588:
1883:. It is tentatively placed within Placodermi as a primitive placoderm, though some paleontologists believe the rationale for the placement is inadequate. The paleontologist
1576:, which have eyes on the top of their head. The orbits for the eyes were extremely small, suggesting the eyes were vestigial and that the phyllolepids may have been blind.
712:. However, the tooth and jaw development were not as closely integrated as in modern gnathostomes. These teeth were likely homologous to the teeth of other gnathostomes.
1296:; because of this, the theory that placoderms are the sister group of chondrichthyans has been replaced by the theory that placoderms are a group of basal gnathostomes.
853:, also from the Gogo Formation, have been found with embryos inside them indicating this group also had live bearing ability. The males reproduced by inserting a long
3183:
Rücklin, M.; Donoghue, P. C. J.; Johanson, Z.; Trinajstic, K.; Marone, F.; Stampanoni, M. (2012). "Development of teeth and jaws in the earliest jawed vertebrates".
6082:
704:
published in 2012 concluded that placoderms (at least this particular genus) likely possessed true teeth contrary to some early studies. The teeth had well defined
5955:
884:. While the claspers in cartilaginous fishes are specialized parts of their paired pelvic fins that have been modified for copulation due to changes in the
5227:
1597:("folded teeth") were lightly armoured placoderms with big heads, big eyes and long bodies. They have a strong but superficial resemblance to modern day
4385:
4087:
Ahlberg, P.E.; Trinajstic, K.; Johanson, Z.; Long, J.A. (2009). "Pelvic claspers confirm chondrichthyan-like internal fertilization in arthrodires".
3723:
Zhu, You-an; Li, Qiang; Lu, Jing; Chen, Yang; Wang, Jianhua; Gai, Zhikun; Zhao, Wenjin; Wei, Guangbiao; Yu, Yilun; Ahlberg, Per E.; Zhu, Min (2022).
819:, Western Australia, of a small female placoderm, about 25 cm (10 in) in length, which died in the process of giving birth to a 6 cm (
5372:
889:
were not attached to the pelvic fins, as are the claspers in fish like sharks, they were much more flexible and could probably be rotated forward.
4194:
Long, J.A.; Trinajstic, K. (2010). "The Late
Devonian Gogo Formation Lagerstatte – Exceptional preservation and Diversity in early Vertebrates".
6166:
6156:
3460:"A Devonian Fish Tale: A New Method of Body Length Estimation Suggests Much Smaller Sizes for Dunkleosteus terrelli (Placodermi: Arthrodira)"
5960:
4425:
3809:
Young, G.C.; Goujet, D.; Lelievre, H. (2001). "Extraocular muscles and cranial segmentation in primitive gnathostomes – fossil evidence".
3501:"A new selenosteid placoderm from the Late Devonian of the eastern Anti-Atlas (Morocco) with preserved body outline and its ecomorphology"
847:"), pushing the date of first viviparity back some 200 million years earlier than had been previously known. Specimens of the arthrodire
782:
whose armor practically anchored them to the sea floor. Some placoderms were herbivorous, such as the Middle to Late
Devonian arthrodire
4268:
Long, J.A.; Trinajstic, K.; Johanson, Z. (2009). "Devonian arthrodire embryos and the origin of internal fertilization in vertebrates".
3592:
4390:
4374:
3897:
1605:, and the living and unrelated holocephalians, most of the ptyctodontids are thought to have lived near the sea bottom and preyed on
1405:, was a true "superpredator" of the latest Devonian period, reaching 3 to as much as 8 metres in length. In contrast, the long-nosed
6141:
4188:
4140:
3844:
1259:
624:. Various groups of placoderms were diverse and abundant during the Devonian, but all placoderms became extinct at the end-Devonian
3360:
Burrow, Carol & Turner, Susan (September 1998). "A review of placoderm scales, and their significance in placoderm phylogeny".
5402:
3612:
Brazeau et al, Fossil evidence for a pharyngeal origin of the vertebrate pectoral girdle, Nature volume 623, pages550–554 (2023)
5220:
1389:
had true teeth in addition to tooth plates). The eye sockets are protected by a bony ring, a feature shared by birds and some
1766:
1243:, in his five volumes on fossil fishes, 1833–1843. In those days, placoderms were thought to be shelled jawless fish akin to
4147:
6161:
5965:
3702:
3318:"Bayesian Morphological Clock Methods Resurrect Placoderm Monophyly and Reveal Rapid Early Evolution in Jawed Vertebrates"
1749:
1609:. On account of their lack of armour, some paleontologists have suggested that the Ptyctodontida were not placoderms, but
816:
811:
Extraordinary evidence of internal fertilization in a placoderm was afforded by the discovery in the Gogo
Formation, near
6171:
5972:
4706:
1369:("jointed neck") were the most diverse and numerically successful of the placoderm orders, occupying roles from giant
5413:
153:
5252:
5213:
3259:
Li, Qiang; Zhu, You-an; Lu, Jing; Chen, Yang; Wang, Jianhua; Peng, Lijian; Wei, Guangbiao; Zhu, Min (August 2021).
1385:, lacked teeth, and used instead the sharpened edges of a bony plate, termed a "tooth plate," as a biting surface (
1070:
The
Silurian fossil record of the placoderms is both literally and figuratively fragmented. Until the discovery of
687:
5407:
5940:
835:
1342:
5950:
4418:
1601:. Their armour was reduced to a pattern of small plates around the head and neck. Like the extinct and related
1539:
suggest it is a basal placoderm closest to the ancestral placoderm. Various Early to Middle
Devonian placoderm
1531:
1498:
1015:
5397:
5379:
5998:
5391:
1953:
1430:
6151:
6146:
6022:
4503:
3959:
2711:
However, the cladogram had changed significantly over the years, and the placoderms are now thought to be
1211:
During the Devonian, placoderms went on to inhabit and dominate almost all known aquatic ecosystems, both
641:
401:
564:, which originated among their ranks. In contrast, one 2016 analysis concluded that Placodermi is likely
6108:
4589:
4148:"New bothriolepid fish from the Late Devonian of Victoria, Australia | The Palaeontological Association"
3633:"A novel specimen-based mid-Paleozoic dataset of antiarch placoderms (the most basal jawed vertebrates)"
1618:
on their skin, while ptyctodontids do not; the armoured plates and scales of holocephalians are made of
1043:
869:, both from the Middle Devonian of Australia, suggesting that the basipterygia were used in copulation.
553:
4762:
5145:
4817:
4789:
698:
predators that dwelled in the middle to upper portions of the water column. A study of the arthrodire
6069:
5993:
5945:
5632:
4796:
4780:
4746:
4728:
4368:
4277:
4232:
4203:
4096:
4041:
3980:
3787:
3736:
3644:
3400:
3272:
3192:
1958:
1801:
1773:
1319:
881:
764:
355:
3829:
1701:
placoderms closely related to the rhenanid placoderms. Superficially, acanthoracids resembled scaly
6136:
5195:
5173:
5075:
4831:
4824:
4411:
3069:
2272:
1843:
1792:
1782:
1761:
1445:
1266:, established the details of placoderm anatomy and identified them as true jawed fishes related to
1188:
1151:
as a transitional form between placoderms and other stem-gnathostomes and crown-group gnathostomes.
1054:
580:
340:
4950:
4378:
2387:
1425:
5681:
5529:
5159:
5052:
4894:
4675:
4608:
4578:
4363:
4301:
4256:
4120:
4065:
3940:
3880:"The ancestral morphotype for the gnathostome pectoral fin revisited and the placoderm condition"
3860:
3768:
3298:
3216:
3096:
2736:
2007:
1623:
1511:
1417:
have been found containing unborn fetuses, indicating that arthrodires gave birth to live young.
873:
840:
512:
372:
148:
4936:
4838:
4629:
4622:
4601:
1584:
894:
6113:
5030:
4664:
3164:
1525:
due to anatomical similarities with the other species found at the same locality. According to
6095:
5855:
5803:
5760:
5754:
5717:
5705:
5675:
5330:
5236:
4810:
4769:
4657:
4615:
4344:
4293:
4248:
4184:
4136:
4112:
4057:
4014:
3996:
3932:
3840:
3760:
3752:
3662:
3621:
Benton, M. J. (2005) Vertebrate Palaeontology, Blackwell, 3rd edition, Figure 3.25 on page 73.
3522:
3481:
3428:
3339:
3290:
3208:
1706:
1178:
1001:
859:
857:
into the female. Elongated basipterygia are also found on the phyllolepid placoderms, such as
445:
6100:
3389:"End-Devonian extinction and a bottleneck in the early evolution of modern jawed vertebrates"
5988:
5899:
5882:
5596:
5511:
5101:
4636:
4334:
4326:
4285:
4240:
4211:
4104:
4049:
4004:
3988:
3971:
3924:
3744:
3652:
3512:
3471:
3418:
3408:
3369:
3329:
3280:
3200:
1884:
1687:
1602:
1569:
1526:
1219:. But this diversity ultimately suffered many casualties during the extinction event at the
1166:
1019:
812:
629:
625:
576:
465:
427:
316:
110:
6087:
6060:
5385:
1638:
1354:
1006:
It was thought for a time that placoderms became extinct due to competition from the first
5868:
5831:
5711:
5662:
5483:
5285:
5138:
4866:
4735:
4223:
Long, J.A.; Trinajstic, K.; Young, G.C.; Senden, T. (2008). "Live birth in the Devonian".
3316:
King, Benedict; Qiao, Tuo; Lee, Michael S. Y.; Zhu, Min; Long, John A. (5 December 2016).
2927:
2716:
2359:
2031:
1522:
1506:
1453:
1305:
989:
899:
779:
673:
537:
469:
416:
348:
331:
223:
3911:
Carr, Robert K.; Johanson, Zerina; Ritchie, Alex (July 2009). "The phyllolepid placoderm
1255:
833: in) offspring and was fossilized with the umbilical cord intact. The fossil, named
4315:"An antiarch placoderm shows that pelvic girdles arose at the root of jawed vertebrates"
4281:
4236:
4215:
4207:
4100:
4045:
3984:
3740:
3648:
3404:
3276:
3237:
3196:
1127:
that combines features of primitive arthrodires with jaw anatomy otherwise only seen in
1030:
The earliest identifiable placoderm fossils are of Chinese origin and date to the early
910:
5843:
5821:
5766:
5748:
5699:
5691:
5668:
5626:
5602:
5347:
4715:
4691:
4684:
4562:
4339:
4314:
4009:
3966:
3423:
3388:
3106:
2940:
2783:
2338:
2212:
2162:
2047:
1896:
1858:
1820:
1733:
1729:
1694:
1682:
1610:
1541:
1519:
1515:
1440:("opposite anus") were the second most successful order of placoderms known, after the
1378:
1279:
1123:
1063:
919:
877:
754:
661:
637:
277:
259:
6130:
5792:
5639:
5614:
5563:
5476:
5358:
5338:
5108:
4964:
4908:
4901:
4853:
4488:
4472:
4305:
4260:
4124:
3864:
3772:
3724:
3302:
3135:
3111:
2883:
2827:
2748:
2571:
2554:
2284:
2228:
2177:
1857:") contains another problematic placoderm of uncertain affinity, known only from the
1850:
1594:
1462:
1399:, were well-armoured fishes with flattened bodies. The largest member of this group,
1370:
1278:; however, as more fossils were found, placoderms were accepted as a sister group of
1240:
1228:
1117:
1078:
984:
971:
939:
700:
645:
608:
603:
572:
453:
300:
265:
205:
73:
4164:
4069:
3944:
1448:, who, after incorrectly identifying the first fossils as being those of an armored
926:
123:
5904:
5783:
5723:
5608:
5541:
5436:
4922:
4880:
4873:
4755:
3632:
3373:
3220:
3131:
2960:
2712:
2588:
2537:
2513:
2489:
2322:
2250:
2108:
1854:
1838:
1809:
1565:
1536:
1481:
1475:
1413:
1401:
1390:
1347:
1326:
1293:
1244:
1157:
1106:
1007:
960:
931:
849:
794:
768:
743:
741:"superpredator", preying upon other predators. Other, smaller arthrodires, such as
732:
705:
678:
665:
633:
620:
565:
557:
525:
449:
281:
252:
141:
133:
4395:
4403:
6054:
5891:
5877:
5813:
5742:
5547:
5502:
5449:
5294:
5166:
5122:
5002:
4995:
4943:
4929:
3101:
3083:
2404:
2124:
1980:
1946:
1907:
1892:
1869:
1861:
1805:
1777:
1643:
1558:
1458:
1314:
1286:
1182:
1101:
1072:
955:
880:. The similarities between the structures has been revealed to be an example of
804:
799:
789:
759:
715:
531:
508:
362:
308:
48:
4313:
Zhu, Min; Yu, Xiaobo; Choo, Brian; Wang, Junqing; Jia, Liantao (23 June 2012).
3748:
3499:
Jobbins, Melina; Rücklin, Martin; Ferrón, Humberto G.; Klug, Christian (2022).
1247:. Some naturalists even suggested that they were shelled invertebrates or even
5909:
5808:
5653:
5590:
5535:
5517:
5316:
5307:
5064:
5009:
4971:
4915:
4887:
4466:
3285:
3260:
3091:
3059:
2944:
2907:
2812:
2437:
2198:
1874:
1816:
1797:
1671:
1553:
1493:
1441:
1407:
1395:
1366:
1359:
1212:
1170:
1090:
1058:
is one of the oldest known placoderms, living in what is now China during the
1039:
963:
936:
915:
865:
844:
775:
749:
738:
722:
691:
614:
592:
588:
561:
542:
504:
480:
409:
248:
239:
195:
93:
58:
6045:
4000:
3756:
3666:
3526:
3517:
3500:
3485:
5863:
5620:
5584:
5489:
5041:
4986:
4957:
4650:
4592:
4565:
4053:
3967:"Osteichthyan-like cranial conditions in an Early Devonian stem gnathostome"
3657:
3413:
3334:
3317:
2761:
2454:
2078:
1940:
1925:
1880:
1827:
1824:
1813:
1790:, the pseudopetalichthids had armour made up of a mosaic of tubercles. Like
1787:
1650:
1606:
1470:
1437:
1309:
1263:
1224:
1197:
1193:
1128:
1059:
1035:
993:
967:
943:
669:
657:
599:
584:
549:
492:
488:
431:
271:
242:
165:
98:
42:
4348:
4330:
4297:
4252:
4116:
4061:
4018:
3936:
3915:: Insights into the evolution of feeding mechanisms in jawed vertebrates".
3764:
3432:
3343:
3294:
3212:
1895:. If this is true, then the holocephalians diverged from sharks before the
1049:
978:
1935:
682:, known from over 100 valid species. The vast majority of placoderms were
17:
6039:
5826:
5771:
5728:
5523:
5322:
5300:
5262:
5152:
5023:
5016:
4803:
4643:
4460:
3261:"A new Silurian fish close to the common ancestor of modern gnathostomes"
2305:
1903:
1899:
1738:
1722:
1702:
1698:
1667:
1662:
1658:
1615:
1598:
1573:
1449:
1374:
1323:
1220:
1216:
1205:
1132:
1031:
885:
784:
771:
683:
520:
516:
484:
442:
438:
324:
185:
114:
88:
83:
68:
63:
53:
4289:
4244:
4108:
3992:
3725:"The oldest complete jawed vertebrates from the early Silurian of China"
3204:
1518:. When it was first discovered in 1980, it was originally regarded as a
1292:
Placoderms also share certain anatomical features only with the jawless
950:
5914:
5454:
5444:
5426:
5115:
3928:
2859:
1919:
1627:
1619:
1466:
1139:
1084:
854:
709:
595:
473:
423:
292:
138:
103:
78:
3476:
3459:
5277:
4454:
3589:"The first vertebrate sexual organs evolved as an extra pair of legs"
1248:
1110:
757:
of Western Australia, had streamlined, bullet-shaped head armor, and
695:
461:
434:
412:
175:
6016:
5205:
3879:
1717:
4165:"New phyllolepids from Victoria and the relationships of the group"
1868:
was a thin fish that, when alive, looked vaguely like an elongated
1479:, the limbs were thick and short, while in advanced forms, such as
5733:
1952:
1934:
1837:
1760:
1716:
1681:
1654:
1637:
1583:
1552:
1492:
1424:
1353:
1341:
1275:
1267:
1162:
1155:
The first officially described Silurian placoderm is an antiarch,
1048:
1011:
977:
925:
909:
714:
579:
of the early Silurian. They eventually outcompeted the previously
500:
496:
3163:
Colbert, Edwin H. (Edwin Harris); Knight, Charles Robert (1951).
6074:
5247:
3837:
Recent Advances in the Origin and Early Radiation of Vertebrates
457:
6020:
5209:
4447:
4407:
4369:
Introduction to the Placodermi Extinct armored fishes with jaws
3631:
Pan, Zhaohui; Niu, Zhibin; Xian, Zumin; Zhu, Min (2023-01-03).
1902:
radiation. Critics of Janvier's position say that aside from a
1042:, as well as other, more primitive, groups. Earlier fossils of
1271:
3965:
Giles, Sam; Friedman, Matt; Brazeau, Martin D. (2015-01-12).
1928:
shows the interrelationships of placoderms according to Carr
1186:, also known from distinctively ornamented plates. In 2022,
602:), producing some of the first and most infamous vertebrate
1161:, which is known from distinctively ornamented plates from
3166:
The dinosaur book: the ruling reptiles and their relatives
1910:, the two groups have little else in common anatomically.
1165:, China. It was originally considered to be from the late
3835:. In Arratia, G.; Wilson, M. V. H.; Cloutier, R. (eds.).
3551:
1308:. There are two further controversial orders: One is the
1304:
Currently, Placodermi are divided into eight recognized
3138:, then modern jawed vertebrates represent extant forms.
2719:
than others. The updated cladogram (Zhu et al., 2016):
1887:, as well as other paleontologists, has suggested that
1736:
and were found throughout the world. The petalichthids
774:, strongly supporting the idea that many, if not most,
4364:
Annetta Markussen-Brown, "Devonian Armoured Fish" 2000
3563:
1204:
appearance and diversity at the very beginning of the
1034:. At that time, they were already differentiated into
523:. 380-million-year-old fossils of three other genera,
4183:
Baltimore: The Johns Hopkins University Press, 1996.
4169:
Proceedings of the Linnean Society of New South Wales
3861:"Fish 380 Million Years Old Found With Unborn Embryo"
1239:
The earliest studies of placoderms were published by
1113:, China, known from an almost complete thoracic armor
5896:
5851:
5818:
5800:
5650:
5572:
5499:
5466:
5354:
5343:
5334:
5327:
5312:
5290:
5273:
6029:
5981:
5933:
5842:
5791:
5782:
5690:
5649:
5571:
5562:
5498:
5465:
5435:
5425:
5261:
5093:
5086:
5063:
5040:
4985:
4852:
4779:
4745:
4714:
4674:
4588:
4561:
4514:
4487:
3898:"Philippe Janvier Tree of Life Contributor Profile"
4181:The Rise of Fishes: 500 Million Years of Evolution
3387:Sallan, Lauren & Coates, Michael (June 2010).
4135:Oxford, New York: Oxford University Press, 1998.
495:), as well as the first vertebrates to have true
3839:. Munchen, Germany: Verlag Dr. Friedrich Pfeil.
571:The first identifiable placoderms appear in the
3830:"Placoderm anatomy and phylogeny: new insights"
3393:Proceedings of the National Academy of Sciences
1879:between its tubercles and the tubercles of the
1514:shield") was a long-snouted placoderm from the
1169:, although later study reconsidered its age at
468:(hence the name), and the rest of the body was
3539:
3355:
3353:
1177:plates are very similar to the early Devonian
5221:
4419:
4196:Annual Review of Earth and Planetary Sciences
3178:
3176:
1461:, particularly with later forms. The pair of
1457:sometimes naked rear portions often becoming
8:
1630:and possibly claspers on the head as well.
6017:
5788:
5568:
5432:
5228:
5214:
5206:
5090:
4858:
4720:
4570:
4558:
4493:
4484:
4444:
4426:
4412:
4404:
1529:, anatomical similarities in the brain of
778:were active swimmers, rather than passive
122:
31:
4338:
4008:
3682:"The Antiarchi from Early Silurian Hunan"
3656:
3516:
3475:
3422:
3412:
3333:
3284:
1473:-like limbs. In primitive forms, such as
1046:placoderms have not yet been discovered.
541:, represent the oldest known examples of
1312:Stensioellida, containing the enigmatic
1192:is described from complete fossils from
949:
3155:
3123:
1705:or small, scaly arthrodires with blunt
1393:. Early arthrodires, such as the genus
1800:. As such, placoderm experts consider
1545:have also been inserted in the order.
730:One of the largest known arthrodires,
3878:Carr, Robert K.; et al. (2010).
3828:Goujet, Daniel; Young, Gavin (2004).
3232:
3230:
3169:. New York: McGraw-Hill. p. 153.
2715:, with some being more closer to the
1891:is not a placoderm, but instead is a
1411:measured just 15 cm. Fossils of
1318:; the other is the equally enigmatic
686:, many of which lived at or near the
628:358.9 million years ago, leaving the
7:
3575:
3564:Long, Trinajstic & Johanson 2009
3445:
4216:10.1146/annurev-earth-040809-152416
3458:Engelman, Russell K. (2023-02-21).
2933:
2923:
2899:
2875:
2851:
2805:
2798:
2776:
2754:
2729:
2722:
2529:
2505:
2429:
2379:
2372:
2365:
2297:
2290:
2265:
2243:
2204:
2155:
2148:
2100:
2093:
2071:
2023:
1999:
1972:
1965:
1906:superficially similar to primitive
3505:Frontiers in Ecology and Evolution
3362:Journal of Vertebrate Paleontology
3238:"Fossil reveals oldest live birth"
792:, such as the gigantic arthrodire
720:External anatomy of the placoderm
656:Many placoderms, particularly the
25:
1697:("spine chests") were a group of
1444:. The order's name was coined by
1260:Swedish Museum of Natural History
942:, was a giant armoured placoderm
767:like that of other fast-swimming
552:, consisting of several distinct
5412:
5406:
5401:
5396:
5390:
5384:
5378:
5371:
5191:
5190:
4705:
3958:
3788:"Introduction to the Placodermi"
3076:
3062:
499:. They were also the first fish
479:Placoderms were among the first
152:
1626:, with the males having pelvic
872:The placoderm claspers are not
708:and were made of both bone and
3374:10.1080/02724634.1999.10011135
1804:to be the sister group of the
1767:Pseudopetalicthys problematica
1076:(and then, the discoveries of
898:showcases that the vertebrate
1:
6167:Taxa named by Frederick McCoy
3591:. 8 June 2014. Archived from
1750:Quasipetalichthys haikouensis
1589:Kimbryanodus williamburyensis
1465:were modified into a pair of
548:Placoderms are thought to be
6157:Llandovery first appearances
5973:List of transitional fossils
914:Evolution and extinction of
464:were covered by articulated
3134:in relation to the rest of
640:survivors who subsequently
6188:
5253:Timeline of fish evolution
3749:10.1038/s41586-022-05136-8
3540:Long & Trinajstic 2010
1917:
999:
918:. The diagram is based on
902:evolved from gill arches.
472:or naked depending on the
6007:
5941:Lists of prehistoric fish
5369:
5243:
5187:
4861:
4723:
4703:
4573:
4557:
4538:
4496:
4483:
4443:
3637:Earth System Science Data
3286:10.1016/j.cub.2021.05.053
2958:
2938:
2931:
2921:
2904:
2897:
2880:
2873:
2856:
2849:
2825:
2810:
2803:
2796:
2781:
2774:
2759:
2752:
2734:
2727:
2585:
2568:
2551:
2534:
2527:
2510:
2503:
2486:
2451:
2434:
2427:
2401:
2384:
2377:
2370:
2363:
2336:
2319:
2302:
2295:
2288:
2270:
2263:
2248:
2241:
2226:
2209:
2202:
2175:
2160:
2153:
2146:
2122:
2105:
2098:
2091:
2076:
2069:
2045:
2028:
2021:
2004:
1997:
1977:
1970:
1452:, mistakenly thought the
935:, among the first of the
836:Materpiscis attenboroughi
378:
371:
236:
231:
149:Scientific classification
147:
130:
121:
34:
6142:Prehistoric fish classes
4146:Long, J. A. (May 1983).
3518:10.3389/fevo.2022.969158
2943:(total group, including
1532:Brindabellaspis stensioi
1499:Brindabellaspis stensioi
1020:end-Devonian extinctions
906:Evolution and extinction
5999:Vertebrate paleontology
4375:"PALAEOZOIC FOSSILS UK"
4054:10.1126/science.aah3764
3658:10.5194/essd-15-41-2023
3414:10.1073/pnas.0914000107
1657:fish") were flattened,
1431:Bothriolepis canadensis
1254:In the late 1920s, Dr.
491:from the first pair of
287:"Maxillate placoderms"
27:Extinct Class of Fishes
5094:"maxillate placoderms"
4331:10.1098/rsbl.2011.1033
3689:Vertebrata PalAsiatica
1962:
1950:
1847:
1770:
1726:
1691:
1647:
1591:
1562:
1502:
1434:
1363:
1351:
1300:Taxonomy and phylogeny
1143:, a close relative of
1067:
997:
975:
947:
923:
727:
690:. Many, primarily the
6109:Paleobiology Database
4398:Dunkleosteus terrelli
3917:Journal of Morphology
3913:Cowralepis mclachlani
3811:Journal of Morphology
3680:Wang Junqing (1991).
3335:10.1093/sysbio/syw107
1956:
1938:
1920:Agnatha § Groups
1841:
1764:
1720:
1685:
1641:
1587:
1556:
1496:
1428:
1357:
1345:
1196:, late Llandovery of
1052:
981:
953:
929:
913:
876:with the claspers in
798:, various members of
718:
6162:Devonian extinctions
5994:Transitional fossils
5633:Pseudopetalichthyida
4781:Macropetalichthyidae
4747:Quasipetalichthyidae
4729:Diandongpetalichthys
4391:Placodermi: Overview
3271:(16): 3613–3620.e2.
1959:Diandongpetalichthys
1802:Pseudopetalichthyida
1774:Pseudopetalichthyida
1757:Pseudopetalichthyida
1666:placoderms, such as
1320:Pseudopetalichthyida
882:convergent evolution
878:cartilaginous fishes
850:Incisoscutum ritchei
723:Coccosteus decipiens
560:to all living jawed
507:, the second set of
356:Pseudopetalichthyida
134:Bothriolepis panderi
6172:Paraphyletic groups
4381:on 17 October 2012.
4290:10.1038/nature07732
4282:2009Natur.457.1124L
4276:(7233): 1124–1127.
4245:10.1038/nature06966
4237:2008Natur.453..650L
4208:2010AREPS..38..255L
4163:Long, J.A. (1984).
4109:10.1038/nature08176
4101:2009Natur.460..888A
4046:2016Sci...354..334Z
3993:10.1038/nature14065
3985:2015Natur.520...82G
3741:2022Natur.609..954Z
3649:2023ESSD...15...41P
3405:2010PNAS..10710131S
3399:(22): 10131–10135.
3277:2021CBio...31E3613L
3240:. BBC. May 28, 2008
3205:10.1038/nature11555
3197:2012Natur.491..748R
3070:Paleontology portal
1864:slates of Germany.
1844:Stensioella heintzi
1793:Stensioella heintzi
1783:Stensioella heintzi
1535:and the brain of a
1446:Edward Drinker Cope
1251:-like vertebrates.
1147:that further links
1099:the basal antiarch
5682:Ischnacanthiformes
5530:Cephalaspidomorphi
5160:Pseudopetalichthys
4676:Weejasperaspididae
4590:Palaeacanthaspidae
3929:10.1002/jmor.10719
3322:Systematic Biology
3097:List of placoderms
2947:as stem lineages)
2737:Cephalaspidomorphi
2008:Pseudopetalichthys
1963:
1951:
1848:
1771:
1727:
1692:
1661:, bottom-dwelling
1648:
1624:sexually dimorphic
1592:
1563:
1559:Phyllolepis orvini
1503:
1435:
1364:
1352:
1068:
998:
976:
948:
924:
841:David Attenborough
728:
392:(from Greek πλάξ (
6124:
6123:
6096:Open Tree of Life
6023:Taxon identifiers
6014:
6013:
5929:
5928:
5925:
5924:
5856:Palaeonisciformes
5804:Onychodontiformes
5761:Petalodontiformes
5755:Iniopterygiformes
5718:Phoebodontiformes
5706:Ctenacanthiformes
5676:Diplacanthiformes
5558:
5557:
5331:Myllokunmingiidae
5237:Evolution of fish
5203:
5202:
5183:
5182:
5133:
5132:
4981:
4980:
4848:
4847:
4811:Macropetalichthys
4770:Quasipetalichthys
4763:Pauropetalichthys
4701:
4700:
4553:
4552:
4549:
4548:
4534:
4533:
4231:(7195): 650–652.
4133:Early Vertebrates
4095:(7257): 888–889.
4040:(6310): 334–336.
3735:(7929): 954–958.
3477:10.3390/d15030318
3191:(7426): 748–751.
3052:
3051:
3043:
3042:
3034:
3033:
3025:
3024:
3016:
3015:
3007:
3006:
2998:
2997:
2989:
2988:
2980:
2979:
2971:
2970:
2838:
2837:
2708:
2707:
2699:
2698:
2690:
2689:
2681:
2680:
2672:
2671:
2663:
2662:
2654:
2653:
2645:
2644:
2636:
2635:
2627:
2626:
2618:
2617:
2609:
2608:
2600:
2599:
2475:
2474:
2466:
2465:
2416:
2415:
2349:
2348:
2188:
2187:
2135:
2134:
2058:
2057:
1002:Evolution of fish
860:Austrophyllolepis
839:(after scientist
387:
386:
227:
16:(Redirected from
6179:
6117:
6116:
6104:
6103:
6091:
6090:
6078:
6077:
6065:
6064:
6063:
6050:
6049:
6048:
6018:
5989:Prehistoric life
5900:Pycnodontiformes
5898:
5883:Acipenseriformes
5853:
5820:
5802:
5789:
5652:
5597:Brindabellaspida
5574:
5569:
5512:Pteraspidomorphi
5501:
5468:
5433:
5417:
5416:
5410:
5405:
5400:
5395:
5394:
5388:
5383:
5382:
5375:
5356:
5345:
5336:
5329:
5314:
5292:
5275:
5230:
5223:
5216:
5207:
5194:
5193:
5146:Holopetalichthys
5102:Bianchengichthys
5091:
4859:
4818:Notopetalichthys
4790:Ellopetalichthys
4721:
4709:
4637:Palaeacanthaspis
4571:
4559:
4494:
4485:
4478:
4477:
4445:
4428:
4421:
4414:
4405:
4382:
4377:. Archived from
4352:
4342:
4309:
4264:
4219:
4176:
4159:
4128:
4080:Other references
4074:
4073:
4029:
4023:
4022:
4012:
3963:
3962:
3955:
3949:
3948:
3908:
3902:
3901:
3894:
3888:
3887:
3875:
3869:
3868:
3857:
3851:
3850:
3834:
3825:
3819:
3818:
3806:
3800:
3799:
3797:
3795:
3783:
3777:
3776:
3720:
3714:
3713:
3711:
3705:. Archived from
3701:
3686:
3677:
3671:
3670:
3660:
3628:
3622:
3619:
3613:
3610:
3604:
3603:
3601:
3600:
3585:
3579:
3573:
3567:
3561:
3555:
3552:Long et al. 2008
3549:
3543:
3537:
3531:
3530:
3520:
3496:
3490:
3489:
3479:
3455:
3449:
3443:
3437:
3436:
3426:
3416:
3384:
3378:
3377:
3357:
3348:
3347:
3337:
3313:
3307:
3306:
3288:
3256:
3250:
3249:
3247:
3245:
3234:
3225:
3224:
3180:
3171:
3170:
3160:
3139:
3128:
3086:
3081:
3080:
3079:
3072:
3067:
3066:
3065:
2934:
2924:
2900:
2876:
2852:
2806:
2799:
2777:
2755:
2730:
2723:
2530:
2506:
2430:
2380:
2373:
2366:
2298:
2291:
2266:
2244:
2205:
2156:
2149:
2101:
2094:
2072:
2024:
2000:
1973:
1966:
1885:Philippe Janvier
1780:, Germany. Like
1688:Palaeacanthaspis
1603:acanthothoracids
1527:Philippe Janvier
1489:Brindabellaspida
1333:Placoderm orders
1235:History of study
968:Middle Famennian
832:
831:
827:
824:
813:Fitzroy Crossing
788:, and some were
626:Hangenberg event
618:and the massive
577:Llandovery epoch
575:during the late
428:prehistoric fish
317:Bianchengichthys
222:
217:
157:
156:
126:
108:
45:
38:Temporal range:
32:
21:
6187:
6186:
6182:
6181:
6180:
6178:
6177:
6176:
6127:
6126:
6125:
6120:
6112:
6107:
6099:
6094:
6086:
6081:
6073:
6068:
6059:
6058:
6053:
6044:
6043:
6038:
6025:
6015:
6010:
6003:
5977:
5921:
5869:Polypteriformes
5838:
5832:Tetrapodomorpha
5778:
5712:Hybodontiformes
5686:
5663:Acanthodiformes
5645:
5554:
5494:
5484:Paraconodontida
5461:
5429:
5421:
5420:
5419:
5411:
5389:
5377:
5367:
5286:Cephalochordate
5257:
5239:
5234:
5204:
5199:
5179:
5139:Brindabellaspis
5129:
5082:
5059:
5036:
4977:
4867:Austroptyctodus
4844:
4797:Epipetalichthys
4775:
4741:
4736:Neopetalichthys
4710:
4697:
4670:
4584:
4545:
4530:
4510:
4479:
4450:
4449:
4439:
4432:
4373:
4360:
4355:
4319:Biology Letters
4312:
4267:
4222:
4193:
4162:
4145:
4086:
4082:
4077:
4031:
4030:
4026:
3979:(7545): 82–85.
3964:
3957:
3956:
3952:
3910:
3909:
3905:
3896:
3895:
3891:
3877:
3876:
3872:
3867:. June 6, 2008.
3859:
3858:
3854:
3847:
3832:
3827:
3826:
3822:
3808:
3807:
3803:
3793:
3791:
3786:Waggoner, Ben.
3785:
3784:
3780:
3722:
3721:
3717:
3709:
3699:
3684:
3679:
3678:
3674:
3630:
3629:
3625:
3620:
3616:
3611:
3607:
3598:
3596:
3587:
3586:
3582:
3574:
3570:
3562:
3558:
3550:
3546:
3538:
3534:
3498:
3497:
3493:
3457:
3456:
3452:
3444:
3440:
3386:
3385:
3381:
3359:
3358:
3351:
3315:
3314:
3310:
3265:Current Biology
3258:
3257:
3253:
3243:
3241:
3236:
3235:
3228:
3182:
3181:
3174:
3162:
3161:
3157:
3153:
3148:
3143:
3142:
3129:
3125:
3120:
3082:
3077:
3075:
3068:
3063:
3061:
3058:
3053:
3044:
3035:
3026:
3017:
3008:
2999:
2990:
2981:
2972:
2928:Eugnathostomata
2839:
2717:Eugnathostomata
2709:
2700:
2691:
2682:
2673:
2664:
2655:
2646:
2637:
2628:
2619:
2610:
2601:
2476:
2467:
2417:
2360:Eubrachythoraci
2350:
2189:
2136:
2059:
2032:Brindabellaspis
1922:
1916:
1836:
1759:
1715:
1680:
1636:
1582:
1551:
1523:acanthothoracid
1507:Brindabellaspis
1491:
1423:
1379:bottom dwellers
1340:
1335:
1302:
1280:chondrichthyans
1237:
1028:
1004:
990:Middle Devonian
908:
900:shoulder girdle
829:
825:
822:
820:
726:
694:, were active,
654:
652:Characteristics
538:Austroptyctodus
466:armoured plates
349:Brindabellaspis
332:Eugnathostomata
221:
215:
151:
117:
107:
106:
101:
96:
91:
86:
81:
76:
71:
66:
61:
56:
51:
41:439–358.9
40:
39:
36:
28:
23:
22:
15:
12:
11:
5:
6185:
6183:
6175:
6174:
6169:
6164:
6159:
6154:
6149:
6144:
6139:
6129:
6128:
6122:
6121:
6119:
6118:
6105:
6092:
6079:
6066:
6051:
6035:
6033:
6027:
6026:
6021:
6012:
6011:
6008:
6005:
6004:
6002:
6001:
5996:
5991:
5985:
5983:
5979:
5978:
5976:
5975:
5970:
5969:
5968:
5963:
5958:
5953:
5948:
5937:
5935:
5931:
5930:
5927:
5926:
5923:
5922:
5920:
5919:
5918:
5917:
5912:
5907:
5902:
5888:
5887:
5886:
5885:
5874:
5873:
5872:
5871:
5860:
5859:
5848:
5846:
5840:
5839:
5837:
5836:
5835:
5834:
5829:
5824:
5822:Porolepiformes
5811:
5806:
5797:
5795:
5786:
5780:
5779:
5777:
5776:
5775:
5774:
5769:
5767:Symmoriiformes
5763:
5757:
5751:
5749:Eugeneodontida
5739:
5738:
5737:
5736:
5731:
5726:
5720:
5714:
5708:
5700:Elasmobranchii
5696:
5694:
5688:
5687:
5685:
5684:
5678:
5672:
5669:Climatiiformes
5665:
5658:
5656:
5647:
5646:
5644:
5643:
5636:
5629:
5627:Acanthothoraci
5623:
5617:
5611:
5605:
5603:Petalichthyida
5599:
5593:
5587:
5580:
5578:
5566:
5560:
5559:
5556:
5555:
5553:
5552:
5551:
5550:
5544:
5538:
5526:
5520:
5514:
5507:
5505:
5496:
5495:
5493:
5492:
5486:
5480:
5477:Protoconodonta
5472:
5470:
5463:
5462:
5460:
5459:
5458:
5457:
5447:
5441:
5439:
5430:
5423:
5422:
5370:
5368:
5366:
5365:
5364:
5363:
5352:
5348:Myllokunmingia
5325:
5320:
5305:
5304:
5303:
5298:
5283:
5282:
5281:
5267:
5265:
5259:
5258:
5256:
5255:
5250:
5244:
5241:
5240:
5235:
5233:
5232:
5225:
5218:
5210:
5201:
5200:
5198:
5188:
5185:
5184:
5181:
5180:
5178:
5177:
5170:
5163:
5156:
5149:
5142:
5134:
5131:
5130:
5128:
5127:
5119:
5112:
5105:
5097:
5095:
5088:
5084:
5083:
5081:
5080:
5079:
5078:
5069:
5067:
5061:
5060:
5058:
5057:
5056:
5055:
5046:
5044:
5038:
5037:
5035:
5034:
5027:
5020:
5013:
5006:
4999:
4991:
4989:
4983:
4982:
4979:
4978:
4976:
4975:
4968:
4961:
4954:
4947:
4940:
4933:
4926:
4919:
4912:
4905:
4898:
4891:
4884:
4877:
4870:
4862:
4856:
4850:
4849:
4846:
4845:
4843:
4842:
4835:
4828:
4821:
4814:
4807:
4800:
4793:
4785:
4783:
4777:
4776:
4774:
4773:
4766:
4759:
4751:
4749:
4743:
4742:
4740:
4739:
4732:
4724:
4718:
4716:Petalichthyida
4712:
4711:
4704:
4702:
4699:
4698:
4696:
4695:
4692:Weejasperaspis
4688:
4685:Murrindalaspis
4680:
4678:
4672:
4671:
4669:
4668:
4661:
4654:
4647:
4640:
4633:
4626:
4619:
4612:
4605:
4597:
4595:
4586:
4585:
4583:
4582:
4574:
4568:
4563:Acanthothoraci
4555:
4554:
4551:
4550:
4547:
4546:
4539:
4536:
4535:
4532:
4531:
4529:
4528:
4527:
4526:
4520:
4518:
4512:
4511:
4509:
4508:
4507:
4506:
4497:
4491:
4481:
4480:
4476:
4475:
4469:
4463:
4457:
4448:
4441:
4440:
4433:
4431:
4430:
4423:
4416:
4408:
4402:
4401:
4396:BBC—report on
4393:
4388:
4383:
4371:
4366:
4359:
4358:External links
4356:
4354:
4353:
4325:(3): 453–456.
4310:
4265:
4220:
4191:
4177:
4160:
4143:
4129:
4083:
4081:
4078:
4076:
4075:
4024:
3950:
3923:(7): 775–804.
3903:
3889:
3870:
3852:
3845:
3820:
3801:
3778:
3715:
3712:on 2013-12-12.
3695:(3): 240–244.
3672:
3623:
3614:
3605:
3580:
3568:
3556:
3544:
3532:
3491:
3450:
3438:
3379:
3368:(2): 204–219.
3349:
3328:(4): 499–516.
3308:
3251:
3226:
3172:
3154:
3152:
3149:
3147:
3144:
3141:
3140:
3122:
3121:
3119:
3116:
3115:
3114:
3109:
3107:Chondrichthyes
3104:
3099:
3094:
3088:
3087:
3073:
3057:
3054:
3050:
3049:
3046:
3045:
3041:
3040:
3037:
3036:
3032:
3031:
3028:
3027:
3023:
3022:
3019:
3018:
3014:
3013:
3010:
3009:
3005:
3004:
3001:
3000:
2996:
2995:
2992:
2991:
2987:
2986:
2983:
2982:
2978:
2977:
2974:
2973:
2969:
2968:
2965:
2964:
2957:
2954:
2953:
2950:
2949:
2941:Chondrichthyes
2937:
2932:
2930:
2922:
2920:
2917:
2916:
2913:
2912:
2903:
2898:
2896:
2893:
2892:
2889:
2888:
2879:
2874:
2872:
2869:
2868:
2865:
2864:
2855:
2850:
2848:
2845:
2844:
2841:
2840:
2836:
2835:
2832:
2831:
2824:
2821:
2820:
2817:
2816:
2809:
2804:
2802:
2797:
2795:
2792:
2791:
2788:
2787:
2784:Petalichthyida
2780:
2775:
2773:
2770:
2769:
2766:
2765:
2758:
2753:
2751:
2745:
2744:
2741:
2740:
2733:
2728:
2726:
2721:
2706:
2705:
2702:
2701:
2697:
2696:
2693:
2692:
2688:
2687:
2684:
2683:
2679:
2678:
2675:
2674:
2670:
2669:
2666:
2665:
2661:
2660:
2657:
2656:
2652:
2651:
2648:
2647:
2643:
2642:
2639:
2638:
2634:
2633:
2630:
2629:
2625:
2624:
2621:
2620:
2616:
2615:
2612:
2611:
2607:
2606:
2603:
2602:
2598:
2597:
2594:
2593:
2584:
2581:
2580:
2577:
2576:
2567:
2564:
2563:
2560:
2559:
2550:
2547:
2546:
2543:
2542:
2533:
2528:
2526:
2523:
2522:
2519:
2518:
2509:
2504:
2502:
2499:
2498:
2495:
2494:
2485:
2482:
2481:
2478:
2477:
2473:
2472:
2469:
2468:
2464:
2463:
2460:
2459:
2450:
2447:
2446:
2443:
2442:
2433:
2428:
2426:
2423:
2422:
2419:
2418:
2414:
2413:
2410:
2409:
2400:
2397:
2396:
2393:
2392:
2383:
2378:
2376:
2371:
2369:
2364:
2362:
2356:
2355:
2352:
2351:
2347:
2346:
2343:
2342:
2339:Buchanosteidae
2335:
2332:
2331:
2328:
2327:
2318:
2315:
2314:
2311:
2310:
2301:
2296:
2294:
2289:
2287:
2281:
2280:
2277:
2276:
2269:
2264:
2262:
2259:
2258:
2255:
2254:
2247:
2242:
2240:
2237:
2236:
2233:
2232:
2225:
2222:
2221:
2218:
2217:
2213:Wuttagoonaspis
2208:
2203:
2201:
2195:
2194:
2191:
2190:
2186:
2185:
2182:
2181:
2174:
2171:
2170:
2167:
2166:
2163:Petalichthyida
2159:
2154:
2152:
2147:
2145:
2142:
2141:
2138:
2137:
2133:
2132:
2129:
2128:
2121:
2118:
2117:
2114:
2113:
2104:
2099:
2097:
2092:
2090:
2087:
2086:
2083:
2082:
2075:
2070:
2068:
2065:
2064:
2061:
2060:
2056:
2055:
2052:
2051:
2048:Acanthothoraci
2044:
2041:
2040:
2037:
2036:
2027:
2022:
2020:
2017:
2016:
2013:
2012:
2003:
1998:
1996:
1990:
1989:
1986:
1985:
1976:
1971:
1969:
1964:
1924:The following
1915:
1912:
1908:holocephalians
1897:Chondrichthyan
1872:, or a skinny
1859:Lower Devonian
1835:
1832:
1821:Acanthothoraci
1758:
1755:
1734:Early Devonian
1730:Petalichthyida
1714:
1713:Petalichthyida
1711:
1695:Acanthothoraci
1679:
1678:Acanthothoraci
1676:
1635:
1632:
1611:holocephalians
1581:
1578:
1550:
1547:
1542:incertae sedis
1520:weejasperaspid
1516:Early Devonian
1490:
1487:
1422:
1419:
1371:apex predators
1339:
1336:
1334:
1331:
1301:
1298:
1236:
1233:
1153:
1152:
1136:
1124:incertae sedis
1121:, a placoderm
1114:
1064:Early Silurian
1027:
1024:
940:apex predators
920:Michael Benton
907:
904:
780:ambush-hunters
755:Gogo Formation
753:, both of the
719:
662:Petalichthyida
653:
650:
638:chondrichthyan
604:apex predators
448:. While their
408:) 'skin') are
385:
384:
383:
382:
376:
375:
369:
368:
367:
366:
358:
352:
344:
336:
335:
334:
328:
320:
312:
304:
296:
285:
278:Acanthothoraci
274:
268:
262:
260:Petalichthyida
256:
245:
234:
233:
229:
228:
213:
209:
208:
203:
199:
198:
193:
189:
188:
183:
179:
178:
173:
169:
168:
163:
159:
158:
145:
144:
128:
127:
119:
118:
102:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
46:
37:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
6184:
6173:
6170:
6168:
6165:
6163:
6160:
6158:
6155:
6153:
6152:Silurian fish
6150:
6148:
6147:Devonian fish
6145:
6143:
6140:
6138:
6135:
6134:
6132:
6115:
6110:
6106:
6102:
6097:
6093:
6089:
6084:
6080:
6076:
6071:
6067:
6062:
6056:
6052:
6047:
6041:
6037:
6036:
6034:
6032:
6028:
6024:
6019:
6006:
6000:
5997:
5995:
5992:
5990:
5987:
5986:
5984:
5980:
5974:
5971:
5967:
5964:
5962:
5959:
5957:
5956:cartilaginous
5954:
5952:
5949:
5947:
5944:
5943:
5942:
5939:
5938:
5936:
5932:
5916:
5913:
5911:
5908:
5906:
5903:
5901:
5895:
5894:
5893:
5890:
5889:
5884:
5881:
5880:
5879:
5876:
5875:
5870:
5867:
5866:
5865:
5862:
5861:
5857:
5850:
5849:
5847:
5845:
5841:
5833:
5830:
5828:
5825:
5823:
5817:
5816:
5815:
5812:
5810:
5807:
5805:
5799:
5798:
5796:
5794:
5790:
5787:
5785:
5781:
5773:
5770:
5768:
5764:
5762:
5758:
5756:
5752:
5750:
5746:
5745:
5744:
5741:
5740:
5735:
5732:
5730:
5727:
5725:
5721:
5719:
5715:
5713:
5709:
5707:
5703:
5702:
5701:
5698:
5697:
5695:
5693:
5692:Cartilaginous
5689:
5683:
5679:
5677:
5673:
5670:
5666:
5664:
5660:
5659:
5657:
5655:
5648:
5641:
5640:Stensioellida
5637:
5634:
5630:
5628:
5624:
5622:
5618:
5616:
5615:Ptyctodontida
5612:
5610:
5606:
5604:
5600:
5598:
5594:
5592:
5588:
5586:
5582:
5581:
5579:
5577:
5570:
5567:
5565:
5561:
5549:
5545:
5543:
5539:
5537:
5533:
5532:
5531:
5527:
5525:
5521:
5519:
5515:
5513:
5509:
5508:
5506:
5504:
5497:
5491:
5487:
5485:
5481:
5478:
5474:
5473:
5471:
5464:
5456:
5453:
5452:
5451:
5448:
5446:
5443:
5442:
5440:
5438:
5434:
5431:
5428:
5424:
5418:
5415:
5409:
5404:
5399:
5393:
5387:
5381:
5374:
5361:
5360:
5359:Zhongxiniscus
5353:
5350:
5349:
5341:
5340:
5339:Haikouichthys
5332:
5326:
5324:
5321:
5319:
5318:
5311:
5310:
5309:
5306:
5302:
5299:
5297:
5296:
5289:
5288:
5287:
5284:
5280:
5279:
5272:
5271:
5270:Basal member
5269:
5268:
5266:
5264:
5260:
5254:
5251:
5249:
5246:
5245:
5242:
5238:
5231:
5226:
5224:
5219:
5217:
5212:
5211:
5208:
5197:
5189:
5186:
5176:
5175:
5174:Xiushanosteus
5171:
5169:
5168:
5164:
5162:
5161:
5157:
5155:
5154:
5150:
5148:
5147:
5143:
5141:
5140:
5136:
5135:
5125:
5124:
5120:
5118:
5117:
5113:
5111:
5110:
5109:Entelognathus
5106:
5104:
5103:
5099:
5098:
5096:
5092:
5089:
5085:
5077:
5073:
5072:
5071:
5070:
5068:
5066:
5062:
5054:
5050:
5049:
5048:
5047:
5045:
5043:
5039:
5033:
5032:
5028:
5026:
5025:
5021:
5019:
5018:
5014:
5012:
5011:
5007:
5005:
5004:
5000:
4998:
4997:
4993:
4992:
4990:
4988:
4984:
4974:
4973:
4969:
4967:
4966:
4965:Rhamphodopsis
4962:
4960:
4959:
4955:
4953:
4952:
4948:
4946:
4945:
4941:
4939:
4938:
4934:
4932:
4931:
4927:
4925:
4924:
4920:
4918:
4917:
4913:
4911:
4910:
4909:Eczematolepis
4906:
4904:
4903:
4902:Destnoporella
4899:
4897:
4896:
4892:
4890:
4889:
4885:
4883:
4882:
4878:
4876:
4875:
4871:
4869:
4868:
4864:
4863:
4860:
4857:
4855:
4854:Ptyctodontida
4851:
4841:
4840:
4836:
4834:
4833:
4832:Tongdzuylepis
4829:
4827:
4826:
4825:Shearsbyaspis
4822:
4820:
4819:
4815:
4813:
4812:
4808:
4806:
4805:
4801:
4799:
4798:
4794:
4792:
4791:
4787:
4786:
4784:
4782:
4778:
4772:
4771:
4767:
4765:
4764:
4760:
4758:
4757:
4753:
4752:
4750:
4748:
4744:
4738:
4737:
4733:
4731:
4730:
4726:
4725:
4722:
4719:
4717:
4713:
4708:
4694:
4693:
4689:
4687:
4686:
4682:
4681:
4679:
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4524:
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4513:
4505:
4504:Gnathostomata
4501:
4500:
4499:
4498:
4495:
4492:
4490:
4489:Gnathostomata
4486:
4482:
4474:
4473:Gnathostomata
4471:Infraphylum:
4470:
4468:
4464:
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4458:
4456:
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4230:
4226:
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4213:
4209:
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4197:
4192:
4190:
4189:0-8018-5438-5
4186:
4182:
4178:
4174:
4170:
4166:
4161:
4158:(2): 295–320.
4157:
4153:
4152:Palaeontology
4149:
4144:
4142:
4141:0-19-854047-7
4138:
4134:
4130:
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3865:Science Daily
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3846:3-89937-052-X
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3595:on 2016-12-20
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3186:
3179:
3177:
3173:
3168:
3167:
3159:
3156:
3150:
3145:
3137:
3136:Gnathostomata
3133:
3127:
3124:
3117:
3113:
3112:Entelognathus
3110:
3108:
3105:
3103:
3100:
3098:
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2910:
2909:
2902:
2901:
2895:
2894:
2891:
2890:
2887:
2886:
2885:
2884:Entelognathus
2878:
2877:
2871:
2870:
2867:
2866:
2863:
2862:
2861:
2854:
2853:
2847:
2846:
2843:
2842:
2834:
2833:
2830:
2829:
2828:Ptyctodontida
2823:
2822:
2819:
2818:
2815:
2814:
2808:
2807:
2801:
2800:
2794:
2793:
2790:
2789:
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2785:
2779:
2778:
2772:
2771:
2768:
2767:
2764:
2763:
2757:
2756:
2750:
2749:Gnathostomata
2747:
2746:
2743:
2742:
2739:
2738:
2732:
2731:
2725:
2724:
2720:
2718:
2714:
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2590:
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2579:
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2575:
2574:
2573:
2572:Gorgonichthys
2566:
2565:
2562:
2561:
2558:
2557:
2556:
2555:Erromenosteus
2549:
2548:
2545:
2544:
2541:
2540:
2539:
2532:
2531:
2525:
2524:
2521:
2520:
2517:
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2508:
2507:
2501:
2500:
2497:
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2317:
2316:
2313:
2312:
2309:
2308:
2307:
2300:
2299:
2293:
2292:
2286:
2285:Brachythoraci
2283:
2282:
2279:
2278:
2275:
2274:
2273:Phlyctaeniida
2268:
2267:
2261:
2260:
2257:
2256:
2253:
2252:
2246:
2245:
2239:
2238:
2235:
2234:
2231:
2230:
2229:Actinolepidae
2224:
2223:
2220:
2219:
2216:
2215:
2214:
2207:
2206:
2200:
2197:
2196:
2193:
2192:
2184:
2183:
2180:
2179:
2178:Ptyctodontida
2173:
2172:
2169:
2168:
2165:
2164:
2158:
2157:
2151:
2150:
2144:
2143:
2140:
2139:
2131:
2130:
2127:
2126:
2120:
2119:
2116:
2115:
2112:
2111:
2110:
2103:
2102:
2096:
2095:
2089:
2088:
2085:
2084:
2081:
2080:
2074:
2073:
2067:
2066:
2063:
2062:
2054:
2053:
2050:
2049:
2043:
2042:
2039:
2038:
2035:
2034:
2033:
2026:
2025:
2019:
2018:
2015:
2014:
2011:
2010:
2009:
2002:
2001:
1995:
1992:
1991:
1988:
1987:
1984:
1983:
1982:
1975:
1974:
1968:
1967:
1961:
1960:
1955:
1949:
1948:
1943:
1942:
1937:
1933:
1931:
1927:
1921:
1913:
1911:
1909:
1905:
1901:
1898:
1894:
1893:holocephalian
1890:
1886:
1882:
1877:
1876:
1871:
1867:
1863:
1860:
1856:
1852:
1851:Stensioellida
1846:
1845:
1840:
1834:Stensioellida
1833:
1831:
1829:
1826:
1822:
1818:
1815:
1811:
1807:
1803:
1799:
1795:
1794:
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1784:
1779:
1775:
1769:
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1731:
1725:
1724:
1719:
1712:
1710:
1708:
1704:
1700:
1699:chimaera-like
1696:
1690:
1689:
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1677:
1675:
1673:
1669:
1664:
1660:
1656:
1652:
1646:
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1633:
1631:
1629:
1625:
1621:
1617:
1612:
1608:
1604:
1600:
1596:
1595:Ptyctodontida
1590:
1586:
1580:Ptyctodontida
1579:
1577:
1575:
1571:
1567:
1561:
1560:
1555:
1548:
1546:
1544:
1543:
1538:
1534:
1533:
1528:
1524:
1521:
1517:
1513:
1512:Brindabella's
1509:
1508:
1501:
1500:
1495:
1488:
1486:
1484:
1483:
1478:
1477:
1472:
1468:
1464:
1463:pectoral fins
1460:
1455:
1451:
1447:
1443:
1439:
1433:
1432:
1427:
1420:
1418:
1416:
1415:
1410:
1409:
1404:
1403:
1398:
1397:
1392:
1388:
1387:Compagopiscis
1384:
1383:Compagopiscis
1380:
1376:
1372:
1368:
1362:
1361:
1356:
1350:
1349:
1344:
1337:
1332:
1330:
1328:
1325:
1321:
1317:
1316:
1311:
1307:
1299:
1297:
1295:
1294:osteostracans
1290:
1288:
1283:
1281:
1277:
1273:
1269:
1265:
1261:
1257:
1252:
1250:
1246:
1242:
1241:Louis Agassiz
1234:
1232:
1230:
1229:Carboniferous
1226:
1222:
1218:
1214:
1209:
1207:
1201:
1199:
1195:
1191:
1190:
1189:Xiushanosteus
1185:
1184:
1180:
1176:
1172:
1168:
1164:
1160:
1159:
1150:
1149:Entelognathus
1146:
1145:Entelognathus
1142:
1141:
1137:
1134:
1130:
1126:
1125:
1120:
1119:
1118:Entelognathus
1115:
1112:
1108:
1104:
1103:
1098:
1097:
1096:
1093:
1092:
1087:
1086:
1081:
1080:
1079:Entelognathus
1075:
1074:
1065:
1062:stage of the
1061:
1057:
1056:
1055:Xiushanosteus
1051:
1047:
1045:
1041:
1037:
1033:
1026:Fossil record
1025:
1023:
1021:
1017:
1016:Late Devonian
1013:
1009:
1003:
995:
991:
987:
986:
985:Eczematolepis
982:Fin spine of
980:
973:
972:Late Devonian
969:
965:
962:
958:
957:
952:
945:
941:
938:
934:
933:
928:
921:
917:
912:
905:
903:
901:
897:
896:
890:
887:
883:
879:
875:
870:
868:
867:
862:
861:
856:
852:
851:
846:
842:
838:
837:
818:
814:
809:
807:
806:
801:
797:
796:
791:
787:
786:
781:
777:
773:
770:
766:
762:
761:
756:
752:
751:
746:
745:
740:
736:
734:
725:
724:
717:
713:
711:
707:
706:pulp cavities
703:
702:
701:Compagopiscis
697:
693:
689:
685:
681:
680:
675:
671:
667:
663:
659:
651:
649:
647:
646:Carboniferous
643:
639:
635:
632:open for the
631:
627:
623:
622:
617:
616:
611:
610:
609:Eastmanosteus
605:
601:
597:
594:
590:
586:
582:
578:
574:
573:fossil record
569:
567:
563:
559:
555:
551:
546:
544:
540:
539:
534:
533:
528:
527:
522:
518:
515:precursor to
514:
510:
506:
502:
498:
494:
490:
486:
482:
477:
475:
471:
467:
463:
459:
455:
454:cartilaginous
451:
450:endoskeletons
447:
444:
440:
436:
433:
429:
425:
421:
418:
414:
411:
407:
404:' and δέρμα (
403:
399:
395:
391:
380:
379:
377:
374:
370:
365:
364:
359:
357:
353:
351:
350:
345:
343:
342:
341:Xiushanosteus
337:
333:
329:
327:
326:
321:
319:
318:
313:
311:
310:
305:
303:
302:
301:Entelognathus
297:
295:
294:
289:
288:
286:
283:
279:
275:
273:
269:
267:
266:Ptyctodontida
263:
261:
257:
254:
250:
246:
244:
241:
238:
237:
235:
230:
225:
220:
214:
211:
210:
207:
206:Gnathostomata
204:
202:Infraphylum:
201:
200:
197:
194:
191:
190:
187:
184:
181:
180:
177:
174:
171:
170:
167:
164:
161:
160:
155:
150:
146:
143:
142:pectoral fins
140:
136:
135:
129:
125:
120:
116:
112:
105:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
44:
33:
30:
19:
6030:
5946:spiny sharks
5905:Halecomorphi
5724:Xenacanthida
5609:Phyllolepida
5575:
5542:Pituriaspida
5503:Ostracoderms
5437:Cyclostomata
5427:Jawless fish
5376:
5357:
5346:
5337:
5315:
5293:
5276:
5172:
5165:
5158:
5151:
5144:
5137:
5121:
5114:
5107:
5100:
5029:
5022:
5015:
5008:
5001:
4994:
4970:
4963:
4956:
4951:Ptyctodopsis
4949:
4942:
4935:
4928:
4923:Kimbryanodus
4921:
4914:
4907:
4900:
4893:
4886:
4881:Chelyophorus
4879:
4874:Campbellodus
4872:
4865:
4837:
4830:
4823:
4816:
4809:
4802:
4795:
4788:
4768:
4761:
4756:Eurycaraspis
4754:
4734:
4727:
4690:
4683:
4663:
4656:
4649:
4642:
4635:
4628:
4621:
4614:
4607:
4600:
4577:
4541:
4540:
4515:
4435:
4397:
4379:the original
4322:
4318:
4273:
4269:
4228:
4224:
4199:
4195:
4180:
4172:
4168:
4155:
4151:
4132:
4131:Janvier, P.
4092:
4088:
4037:
4033:
4027:
3976:
3970:
3953:
3920:
3916:
3912:
3906:
3892:
3883:
3873:
3855:
3836:
3823:
3814:
3810:
3804:
3792:. Retrieved
3781:
3732:
3728:
3718:
3707:the original
3692:
3688:
3675:
3643:(1): 41–51.
3640:
3636:
3626:
3617:
3608:
3597:. Retrieved
3593:the original
3583:
3571:
3559:
3547:
3535:
3508:
3504:
3494:
3467:
3463:
3453:
3441:
3396:
3392:
3382:
3365:
3361:
3325:
3321:
3311:
3268:
3264:
3254:
3242:. Retrieved
3188:
3184:
3165:
3158:
3132:paraphyletic
3126:
2961:Osteichthyes
2959:
2939:
2906:
2905:
2882:
2881:
2858:
2857:
2826:
2811:
2782:
2760:
2735:
2713:paraphyletic
2710:
2589:Titanichthys
2587:
2586:
2570:
2569:
2553:
2552:
2538:Brachyosteus
2536:
2535:
2514:Dunkleosteus
2512:
2511:
2490:Plourdosteus
2488:
2487:
2453:
2452:
2436:
2435:
2403:
2402:
2388:Pholidosteus
2386:
2385:
2337:
2323:Antineosteus
2321:
2320:
2304:
2303:
2271:
2251:Phyllolepida
2249:
2227:
2211:
2210:
2176:
2161:
2123:
2109:Yunnanolepis
2107:
2106:
2077:
2046:
2030:
2029:
2006:
2005:
1993:
1979:
1978:
1957:
1945:
1939:
1929:
1923:
1888:
1873:
1865:
1849:
1842:
1810:Phyllolepida
1791:
1781:
1772:
1765:
1748:
1743:
1737:
1728:
1721:
1693:
1686:
1649:
1642:
1593:
1588:
1566:Phyllolepida
1564:
1557:
1549:Phyllolepida
1540:
1537:jawless fish
1530:
1505:
1504:
1497:
1482:Bothriolepis
1480:
1476:Yunnanolepis
1474:
1436:
1429:
1414:Incisoscutum
1412:
1406:
1402:Dunkleosteus
1400:
1394:
1391:ichthyosaurs
1386:
1382:
1365:
1358:
1348:Dunkleosteus
1346:
1327:lagerstatten
1313:
1303:
1291:
1284:
1256:Erik Stensiö
1253:
1245:ostracoderms
1238:
1210:
1202:
1187:
1181:
1179:yunnanolepid
1175:Shimenolepis
1174:
1158:Shimenolepis
1156:
1154:
1148:
1144:
1138:
1122:
1116:
1107:Ludlow epoch
1100:
1089:
1083:
1077:
1071:
1069:
1053:
1029:
1005:
983:
954:
932:Dunkleosteus
930:
893:
891:
871:
864:
858:
848:
834:
810:
803:
795:Titanichthys
793:
790:planktivores
783:
758:
748:
744:Fallacosteus
742:
733:Dunkleosteus
731:
729:
721:
699:
679:Bothriolepis
677:
666:Phyllolepida
655:
634:osteichthyan
621:Dunkleosteus
619:
613:
607:
570:
566:monophyletic
550:paraphyletic
547:
536:
530:
526:Incisoscutum
524:
478:
419:
405:
397:
393:
389:
388:
381:Placodermata
361:
347:
339:
323:
315:
307:
299:
291:
282:paraphyletic
253:Phyllolepida
218:
139:caliper-like
137:showing its
132:
29:
6055:Wikispecies
5966:lobe-finned
5892:Neopterygii
5878:Chondrostei
5814:Rhipidistia
5793:Lobe-finned
5743:Holocephali
5548:Osteostraci
5450:Hyperoartia
5295:Cathaymyrus
5263:Forerunners
5167:Stensioella
5123:Silurolepis
5003:Bolivosteus
4996:Asterosteus
4944:Palaeomylus
4930:Materpiscis
4895:Denisonodus
4609:Dobrowlania
4579:Hagiangella
4465:Subphylum:
4202:: 255–279.
4179:Long, J.A.
3102:Ostracoderm
3084:Fish portal
2405:Tapinosteus
2125:Euantiarcha
1981:Stensioella
1947:Pterichthys
1889:Stensioella
1866:Stensioella
1806:Arthrodires
1798:ptyctodonts
1672:arthrodires
1644:Asterosteus
1315:Stensioella
1287:gnathostome
1183:Zhanjilepis
1105:, from the
1102:Silurolepis
1073:Silurolepis
1040:arthrodires
988:, from the
956:Amazichthys
892:A study on
805:Heterosteus
800:Homostiidae
776:arthrodires
760:Amazichthys
692:arthrodires
644:during the
589:eurypterids
562:vertebrates
558:sister taxa
532:Materpiscis
509:paired fins
505:pelvic fins
503:to develop
493:gill arches
452:are mainly
437:during the
430:known from
363:Stensioella
309:Silurolepis
192:Subphylum:
6137:Placoderms
6131:Categories
6061:Placodermi
6031:Placodermi
5951:placoderms
5910:Ginglymodi
5844:Ray-finned
5809:Coelacanth
5654:Acanthodii
5591:Arthrodira
5576:Placoderms
5564:Jawed fish
5536:Galeaspida
5518:Thelodonti
5317:Haikouella
5308:Olfactores
5076:Arthrodira
5065:Arthrodira
5010:Gemuendina
4972:Rhynchodus
4937:Meeksiella
4916:Goniosteus
4888:Ctenurella
4839:Wijdeaspis
4630:Kosoraspis
4623:Kolymaspis
4602:Arabosteus
4542:Placodermi
4525:see below↓
4516:Placodermi
4467:Vertebrata
4436:Placodermi
4386:Placoderms
4175:: 263–308.
3599:2014-06-27
3470:(3): 318.
3146:References
3092:Acanthodii
2945:Acanthodii
2908:Janusiscus
2813:Arthrodira
2438:Coccosteus
2199:Arthrodira
1994:Placodermi
1918:See also:
1875:Gemuendina
1853:(" little
1817:trichotomy
1786:, and the
1744:Wijdeaspis
1570:stargazers
1469:-like, or
1442:Arthrodira
1408:Rolfosteus
1396:Arctolepis
1377:-nibbling
1367:Arthrodira
1360:Coccosteus
1338:Arthrodira
1213:freshwater
1171:Ludfordian
1167:Llandovery
1091:Wangolepis
1010:and early
1000:See also:
964:arthrodire
937:vertebrate
916:placoderms
895:Kolymaspis
874:homologous
866:Cowralepis
845:viviparous
765:morphology
750:Rolfosteus
739:vertebrate
615:Dinichthys
600:orthocones
593:cephalopod
585:arthropods
543:live birth
513:homologous
481:jawed fish
420:Placodermi
410:vertebrate
390:Placoderms
251:(includes
249:Arthrodira
219:Placodermi
196:Vertebrata
131:Fossil of
111:Llandovery
18:Placodermi
6009:† extinct
5864:Cladistia
5621:Rhenanida
5585:Antiarchi
5490:Conodonta
5469:Conodonts
5053:Antiarchi
5042:Antiarchi
5031:Ohioaspis
4987:Rhenanida
4958:Ptyctodus
4665:Tlamaspis
4651:Romundina
4453:Kingdom:
4306:205215898
4261:205213348
4125:205217467
4001:1476-4687
3773:252569910
3757:1476-4687
3667:1866-3508
3576:Long 1984
3527:2296-701X
3486:1424-2818
3464:Diversity
3446:Long 1983
3303:235477130
3151:Citations
2762:Antiarchi
2455:Torosteus
2079:Rhenanida
1941:Homostius
1926:cladogram
1914:Cladogram
1881:rhenanids
1828:dichotomy
1825:Rhenanida
1814:Antiarchi
1788:Rhenanida
1703:chimaeras
1668:antiarchs
1663:predators
1651:Rhenanida
1634:Rhenanida
1607:shellfish
1599:chimaeras
1471:arthropod
1438:Antiarchi
1421:Antiarchi
1310:monotypic
1264:Stockholm
1258:, at the
1225:Famennian
1217:saltwater
1200:, China.
1198:Chongqing
1194:Telychian
1133:tetrapods
1129:bony fish
1060:Telychian
1036:antiarchs
1008:bony fish
994:Wisconsin
966:from the
886:hox genes
817:Kimberley
772:organisms
688:substrate
684:predators
670:Antiarchi
658:Rhenanida
554:outgroups
521:tetrapods
517:hindlimbs
432:Paleozoic
426:group of
272:Rhenanida
243:Antiarchi
172:Kingdom:
166:Eukaryota
35:Placoderm
6040:Wikidata
5827:Lungfish
5772:Chimaera
5524:Anaspida
5323:Tunicate
5301:Lancelet
5196:Category
5153:Minjinia
5024:Nefudina
5017:Jagorina
4804:Lunaspis
4658:Sudaspis
4644:Radotina
4616:Kimaspis
4461:Chordata
4459:Phylum:
4455:Animalia
4349:22219394
4298:19242474
4253:18509443
4117:19597477
4070:45922669
4062:27846567
4019:25581798
3945:45258255
3937:19215000
3884:Academia
3765:36171378
3703:19733953
3433:20479258
3344:27920231
3295:34146483
3213:23075852
3056:See also
2306:Holonema
1932:(2009):
1904:bodyplan
1900:Devonian
1862:Hunsrück
1819:and the
1778:Hunsrück
1739:Lunaspis
1723:Lunaspis
1707:rostrums
1659:ray-like
1628:claspers
1616:shagreen
1574:flatfish
1454:eye-hole
1450:tunicate
1375:detritus
1324:Hunsruck
1221:Frasnian
1206:Devonian
1032:Silurian
944:predator
785:Holonema
735:terrelli
696:nektonic
642:radiated
606:such as
596:molluscs
581:dominant
511:and the
456:, their
443:Devonian
441:and the
439:Silurian
373:Synonyms
325:Minjinia
186:Chordata
182:Phylum:
176:Animalia
162:Domain:
115:Devonian
6101:5292584
6075:4655826
6046:Q130932
5982:Related
5915:Teleost
5455:Lamprey
5445:Hagfish
5116:Qilinyu
4340:3367742
4278:Bibcode
4233:Bibcode
4204:Bibcode
4097:Bibcode
4042:Bibcode
4034:Science
4010:5536226
3981:Bibcode
3794:18 July
3737:Bibcode
3645:Bibcode
3424:2890420
3401:Bibcode
3273:Bibcode
3244:May 30,
3221:4302415
3193:Bibcode
2860:Qilinyu
1870:ratfish
1855:Stensio
1620:dentine
1467:caliper
1459:sinuous
1140:Qilinyu
1085:Qilinyu
970:of the
961:pelagic
922:, 2005.
855:clasper
828:⁄
769:pelagic
763:, with
710:dentine
583:marine
489:evolved
487:likely
483:(their
474:species
446:periods
435:fossils
424:extinct
415:of the
413:animals
293:Qilinyu
232:Orders
212:Class:
113:– Late
5278:Pikaia
5087:Others
4347:
4337:
4304:
4296:
4270:Nature
4259:
4251:
4225:Nature
4187:
4139:
4123:
4115:
4089:Nature
4068:
4060:
4017:
4007:
3999:
3972:Nature
3943:
3935:
3843:
3817:: 304.
3790:. UCMP
3771:
3763:
3755:
3729:Nature
3700:
3665:
3525:
3484:
3431:
3421:
3342:
3301:
3293:
3219:
3211:
3185:Nature
1930:et al.
1306:orders
1276:sharks
1268:sharks
1249:turtle
1111:Yunnan
1012:sharks
802:, and
668:, and
630:niches
598:(e.g.
591:) and
587:(e.g.
470:scaled
462:thorax
398:plakos
226:, 1848
6114:34249
6083:IRMNG
5934:Lists
5734:Shark
4302:S2CID
4257:S2CID
4121:S2CID
4066:S2CID
3941:S2CID
3833:(PDF)
3769:S2CID
3710:(PDF)
3697:INIST
3685:(PDF)
3299:S2CID
3217:S2CID
3118:Notes
1655:Rhine
1163:Hunan
1044:basal
674:genus
501:clade
497:teeth
422:, an
417:class
406:derma
402:plate
224:McCoy
109:Late
6088:1495
5961:bony
5784:Bony
5248:Fish
5074:see
5051:see
4502:see
4345:PMID
4294:PMID
4249:PMID
4185:ISBN
4137:ISBN
4113:PMID
4058:PMID
4015:PMID
3997:ISSN
3933:PMID
3841:ISBN
3796:2011
3761:PMID
3753:ISSN
3663:ISSN
3523:ISSN
3482:ISSN
3429:PMID
3340:PMID
3291:PMID
3246:2008
3209:PMID
1944:and
1742:and
1670:and
1215:and
1131:and
1082:and
1038:and
1018:and
959:, a
863:and
747:and
636:and
535:and
485:jaws
460:and
458:head
394:plax
49:PreꞒ
6070:EoL
5729:Ray
5333:? (
4335:PMC
4327:doi
4286:doi
4274:457
4241:doi
4229:453
4212:doi
4173:107
4105:doi
4093:460
4050:doi
4038:354
4005:PMC
3989:doi
3977:520
3925:doi
3921:270
3815:248
3745:doi
3733:609
3653:doi
3513:doi
3472:doi
3419:PMC
3409:doi
3397:107
3370:doi
3330:doi
3281:doi
3201:doi
3189:491
3130:If
1572:or
1373:to
1272:wax
1262:in
1109:of
992:of
556:or
519:in
400:) '
6133::
6111::
6098::
6085::
6072::
6057::
6042::
5667:†"
5342:,
4343:.
4333:.
4321:.
4317:.
4300:.
4292:.
4284:.
4272:.
4255:.
4247:.
4239:.
4227:.
4210:.
4200:38
4198:.
4171:.
4167:.
4156:26
4154:.
4150:.
4119:.
4111:.
4103:.
4091:.
4064:.
4056:.
4048:.
4036:.
4013:.
4003:.
3995:.
3987:.
3975:.
3969:.
3939:.
3931:.
3919:.
3882:.
3863:.
3813:.
3767:.
3759:.
3751:.
3743:.
3731:.
3727:.
3693:21
3691:.
3687:.
3661:.
3651:.
3641:15
3639:.
3635:.
3521:.
3511:.
3509:10
3507:.
3503:.
3480:.
3468:15
3466:.
3462:.
3427:.
3417:.
3407:.
3395:.
3391:.
3366:19
3364:.
3352:^
3338:.
3326:66
3324:.
3320:.
3297:.
3289:.
3279:.
3269:31
3267:.
3263:.
3229:^
3215:.
3207:.
3199:.
3187:.
3175:^
1830:.
1823:+
1812:+
1808:+
1653:("
1510:("
1329:.
1282:.
1231:.
1208:.
1173:.
1066:.
1022:.
815:,
808:.
676:,
664:,
660:,
648:.
612:,
568:.
545:.
529:,
476:.
396:,
360:?†
354:?†
99:Pg
43:Ma
5897:†
5858:"
5854:"
5852:†
5819:†
5801:†
5765:†
5759:†
5753:†
5747:†
5722:†
5716:†
5710:†
5704:†
5680:†
5674:†
5671:"
5661:†
5651:†
5642:?
5638:†
5635:?
5631:†
5625:†
5619:†
5613:†
5607:†
5601:†
5595:†
5589:†
5583:†
5573:†
5546:†
5540:†
5534:†
5528:†
5522:†
5516:†
5510:†
5500:†
5488:†
5482:†
5479:?
5475:†
5467:†
5362:?
5355:†
5351:)
5344:†
5335:†
5328:†
5313:†
5291:†
5274:†
5229:e
5222:t
5215:v
5126:?
4593:*
4566:*
4438:"
4434:"
4427:e
4420:t
4413:v
4351:.
4329::
4323:8
4308:.
4288::
4280::
4263:.
4243::
4235::
4218:.
4214::
4206::
4127:.
4107::
4099::
4072:.
4052::
4044::
4021:.
3991::
3983::
3947:.
3927::
3900:.
3886:.
3849:.
3798:.
3775:.
3747::
3739::
3669:.
3655::
3647::
3602:.
3578:.
3566:.
3554:.
3542:.
3529:.
3515::
3488:.
3474::
3448:.
3435:.
3411::
3403::
3376:.
3372::
3346:.
3332::
3305:.
3283::
3275::
3248:.
3223:.
3203::
3195::
1223:–
1135:.
996:.
974:.
946:.
830:2
826:1
823:+
821:2
346:†
338:†
330:?
322:†
314:†
306:†
298:†
290:†
284:)
280:(
276:†
270:†
264:†
258:†
255:)
247:†
240:†
216:†
104:N
94:K
89:J
84:T
79:P
74:C
69:D
64:S
59:O
54:Ꞓ
20:)
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