1698:. 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.
5362:
5392:
1332:
5403:
3949:
1415:
1751:
3053:
1574:
1943:
5387:
143:
4696:
5369:
5397:
1628:
1344:
5381:
900:
1672:
5181:
3067:
916:
113:
1828:
705:
1543:
1483:
5375:
1039:
968:
1557:("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
1925:
940:
1707:
1663:. 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.
1785:, 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
1003:, 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
877:
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
1611:, 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
1867:
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
1192:
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
1445:
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,
1263:. 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
1654:
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"
1077:), 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,
3670:
1742:. 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.
1474:, 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.
1735:
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,
1083:
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:
3695:
661:, 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
832:), 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 ("
726:, 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
1721:("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
1602:
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
1765:("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
1274:
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
1311:. 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
1259:. 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
1216:
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
1278:
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.
1370:. 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
4021:
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".
3577:
1872:. It is tentatively placed within Placodermi as a primitive placoderm, though some paleontologists believe the rationale for the placement is inadequate. The paleontologist
1565:, 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.
701:. 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.
1285:; 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.
842:, 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
3172:
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".
693:
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
3685:
873:. 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
4578:
4751:
1586:("folded teeth") were lightly armoured placoderms with big heads, big eyes and long bodies. They have a strong but superficial resemblance to modern day
5134:
4806:
4778:
4785:
4374:
4076:
Ahlberg, P.E.; Trinajstic, K.; Johanson, Z.; Long, J.A. (2009). "Pelvic claspers confirm chondrichthyan-like internal fertilization in arthrodires".
3712:
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).
808:, 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 (
878:
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.
6071:
5162:
4820:
4813:
4183:
Long, J.A.; Trinajstic, K. (2010). "The Late
Devonian Gogo Formation Lagerstatte – Exceptional preservation and Diversity in early Vertebrates".
2261:
1177:
1043:
329:
4939:
2376:
5944:
4883:
4597:
4567:
3449:"A Devonian Fish Tale: A New Method of Body Length Estimation Suggests Much Smaller Sizes for Dunkleosteus terrelli (Placodermi: Arthrodira)"
4925:
4827:
4618:
4611:
4590:
883:
3798:
Young, G.C.; Goujet, D.; Lelievre, H. (2001). "Extraocular muscles and cranial segmentation in primitive gnathostomes – fossil evidence".
3490:"A new selenosteid placoderm from the Late Devonian of the eastern Anti-Atlas (Morocco) with preserved body outline and its ecomorphology"
5216:
5019:
4653:
836:"), pushing the date of first viviparity back some 200 million years earlier than had been previously known. Specimens of the arthrodire
771:
whose armor practically anchored them to the sea floor. Some placoderms were herbivorous, such as the Middle to Late
Devonian arthrodire
4257:
Long, J.A.; Trinajstic, K.; Johanson, Z. (2009). "Devonian arthrodire embryos and the origin of internal fertilization in vertebrates".
3581:
4646:
4604:
4379:
4363:
3886:
1594:, and the living and unrelated holocephalians, most of the ptyctodontids are thought to have lived near the sea bottom and preyed on
1394:, was a true "superpredator" of the latest Devonian period, reaching 3 to as much as 8 metres in length. In contrast, the long-nosed
613:. Various groups of placoderms were diverse and abundant during the Devonian, but all placoderms became extinct at the end-Devonian
3349:
Burrow, Carol & Turner, Susan (September 1998). "A review of placoderm scales, and their significance in placoderm phylogeny".
3601:
Brazeau et al, Fossil evidence for a pharyngeal origin of the vertebrate pectoral girdle, Nature volume 623, pages550–554 (2023)
6155:
1378:
had true teeth in addition to tooth plates). The eye sockets are protected by a bony ring, a feature shared by birds and some
6145:
1232:, in his five volumes on fossil fishes, 1833–1843. In those days, placoderms were thought to be shelled jawless fish akin to
5949:
5361:
4414:
4136:
3691:
3307:"Bayesian Morphological Clock Methods Resurrect Placoderm Monophyly and Reveal Rapid Early Evolution in Jawed Vertebrates"
1598:. On account of their lack of armour, some paleontologists have suggested that the Ptyctodontida were not placoderms, but
800:
Extraordinary evidence of internal fertilization in a placoderm was afforded by the discovery in the Gogo
Formation, near
1358:("jointed neck") were the most diverse and numerically successful of the placoderm orders, occupying roles from giant
6130:
4177:
4129:
3833:
1248:
3248:
Li, Qiang; Zhu, You-an; Lu, Jing; Chen, Yang; Wang, Jianhua; Peng, Lijian; Wei, Guangbiao; Zhu, Min (August 2021).
1374:, lacked teeth, and used instead the sharpened edges of a bony plate, termed a "tooth plate," as a biting surface (
1059:
The
Silurian fossil record of the placoderms is both literally and figuratively fragmented. Until the discovery of
5209:
4437:
1755:
1590:. Their armour was reduced to a pattern of small plates around the head and neck. Like the extinct and related
1528:
suggest it is a basal placoderm closest to the ancestral placoderm. Various Early to Middle
Devonian placoderm
5391:
6150:
5954:
1738:
805:
6160:
5961:
2700:
However, the cladogram had changed significantly over the years, and the placoderms are now thought to be
1200:
During the Devonian, placoderms went on to inhabit and dominate almost all known aquatic ecosystems, both
553:, which originated among their ranks. In contrast, one 2016 analysis concluded that Placodermi is likely
4695:
4137:"New bothriolepid fish from the Late Devonian of Victoria, Australia | The Palaeontological Association"
3622:"A novel specimen-based mid-Paleozoic dataset of antiarch placoderms (the most basal jawed vertebrates)"
1607:
on their skin, while ptyctodontids do not; the armoured plates and scales of holocephalians are made of
858:, both from the Middle Devonian of Australia, suggesting that the basipterygia were used in copulation.
5241:
5202:
676:
142:
687:
predators that dwelled in the middle to upper portions of the water column. A study of the arthrodire
5929:
4357:
3776:
824:
5396:
3948:
3818:
1690:
placoderms closely related to the rhenanid placoderms. Superficially, acanthoracids resembled scaly
5939:
5402:
4407:
1520:
1487:
1255:, established the details of placoderm anatomy and identified them as true jawed fishes related to
1140:
as a transitional form between placoderms and other stem-gnathostomes and crown-group gnathostomes.
1004:
4367:
5987:
4352:
3869:"The ancestral morphotype for the gnathostome pectoral fin revisited and the placoderm condition"
3849:
1419:
1406:
have been found containing unborn fetuses, indicating that arthrodires gave birth to live young.
6140:
6135:
6102:
6011:
5380:
4492:
3153:
1514:
due to anatomical similarities with the other species found at the same locality. According to
630:
390:
5386:
5368:
3610:
Benton, M. J. (2005) Vertebrate Palaeontology, Blackwell, 3rd edition, Figure 3.25 on page 73.
846:
into the female. Elongated basipterygia are also found on the phyllolepid placoderms, such as
6097:
6089:
3378:"End-Devonian extinction and a bottleneck in the early evolution of modern jawed vertebrates"
1032:
542:
1331:
1208:. But this diversity ultimately suffered many casualties during the extinction event at the
6076:
6058:
6049:
5982:
5934:
5621:
4769:
4735:
4717:
4266:
4221:
4192:
4085:
4030:
3969:
3725:
3633:
3389:
3261:
3181:
1947:
1790:
1762:
1308:
995:
It was thought for a time that placoderms became extinct due to competition from the first
870:
753:
344:
4212:
Long, J.A.; Trinajstic, K.; Young, G.C.; Senden, T. (2008). "Live birth in the Devonian".
3305:
King, Benedict; Qiao, Tuo; Lee, Michael S. Y.; Zhu, Min; Long, John A. (5 December 2016).
8:
6125:
5184:
5064:
4400:
3900:
Carr, Robert K.; Johanson, Zerina; Ritchie, Alex (July 2009). "The phyllolepid placoderm
3058:
1942:
1832:
1781:
1771:
1434:
822: in) offspring and was fossilized with the umbilical cord intact. The fossil, named
569:
4304:"An antiarch placoderm shows that pelvic girdles arose at the root of jawed vertebrates"
4270:
4225:
4204:
4196:
4089:
4034:
3973:
3729:
3637:
3393:
3265:
3226:
3185:
1116:
that combines features of primitive arthrodires with jaw anatomy otherwise only seen in
1019:
The earliest identifiable placoderm fossils are of Chinese origin and date to the early
5670:
5518:
5148:
5041:
4664:
4328:
4303:
4290:
4245:
4109:
4054:
3998:
3955:
3929:
3757:
3412:
3377:
3287:
3205:
3085:
2725:
1996:
1612:
1500:
1429:("opposite anus") were the second most successful order of placoderms known, after the
862:
829:
501:
361:
137:
6084:
5844:
5792:
5749:
5743:
5706:
5694:
5664:
5319:
5225:
4799:
4758:
4333:
4294:
4282:
4249:
4237:
4173:
4125:
4113:
4101:
4046:
4003:
3985:
3921:
3829:
3761:
3749:
3741:
3713:
3651:
3511:
3470:
3417:
3328:
3291:
3279:
3197:
1846:") contains another problematic placoderm of uncertain affinity, known only from the
1695:
1388:, were well-armoured fishes with flattened bodies. The largest member of this group,
1267:; however, as more fossils were found, placoderms were accepted as a sister group of
1167:
990:
848:
434:
4153:
4058:
3933:
1437:, who, after incorrectly identifying the first fossils as being those of an armored
5977:
5888:
5871:
5585:
5500:
5090:
4625:
4323:
4315:
4274:
4229:
4200:
4093:
4038:
3993:
3977:
3960:
3913:
3733:
3641:
3621:
3501:
3460:
3407:
3397:
3362:
3358:
3318:
3269:
3209:
3189:
1873:
1676:
1591:
1558:
1515:
1155:
1008:
801:
730:"superpredator", preying upon other predators. Other, smaller arthrodires, such as
618:
614:
565:
454:
416:
305:
99:
4384:
1750:
5857:
5820:
5700:
5651:
5472:
5274:
5127:
4855:
4724:
4392:
2916:
2705:
2348:
2020:
1511:
1495:
1442:
1414:
1294:
978:
888:
768:
662:
526:
458:
405:
337:
320:
212:
5374:
1805:
1573:
1447:
1244:
869:. The similarities between the structures has been revealed to be an example of
5832:
5810:
5755:
5737:
5688:
5680:
5657:
5615:
5591:
5336:
4704:
4680:
4673:
4551:
4302:
Zhu, Min; Yu, Xiaobo; Choo, Brian; Wang, Junqing; Jia, Liantao (23 June 2012).
3737:
3488:
Jobbins, Melina; Rücklin, Martin; Ferrón, Humberto G.; Klug, Christian (2022).
3095:
2929:
2772:
2327:
2201:
2151:
2036:
1885:
1847:
1809:
1722:
1718:
1683:
1599:
1530:
1508:
1504:
1367:
1268:
1236:. Some naturalists even suggested that they were shelled invertebrates or even
1112:
1052:
908:
866:
743:
650:
626:
266:
248:
3274:
3249:
1047:
is one of the oldest known placoderms, living in what is now China during the
6119:
6034:
5781:
5628:
5603:
5552:
5465:
5347:
5327:
5097:
4953:
4897:
4890:
4842:
4477:
4461:
3989:
3853:
3745:
3655:
3515:
3506:
3489:
3474:
3124:
3100:
2872:
2816:
2737:
2560:
2543:
2273:
2217:
2166:
1839:
1583:
1451:
1359:
1229:
1217:
1106:
1067:
973:
960:
928:
689:
634:
597:
592:
561:
442:
289:
254:
194:
62:
4042:
3956:"Osteichthyan-like cranial conditions in an Early Devonian stem gnathostome"
3646:
3402:
3323:
3306:
1779:, the pseudopetalichthids had armour made up of a mosaic of tubercles. Like
1627:
1343:
5893:
5772:
5712:
5597:
5530:
5425:
4911:
4869:
4862:
4744:
4337:
4319:
4286:
4241:
4105:
4050:
4007:
3925:
3904:: Insights into the evolution of feeding mechanisms in jawed vertebrates".
3753:
3421:
3332:
3283:
3201:
3120:
2949:
2701:
2577:
2526:
2502:
2478:
2311:
2239:
2097:
1884:. If this is true, then the holocephalians diverged from sharks before the
1843:
1798:
1554:
1525:
1470:
1464:
1402:
1390:
1379:
1336:
1315:
1282:
1233:
1146:
1095:
996:
949:
920:
838:
783:
757:
732:
721:
694:
667:
654:
622:
609:
554:
546:
514:
438:
270:
241:
130:
122:
899:
671:, known from over 100 valid species. The vast majority of placoderms were
6043:
5880:
5866:
5802:
5731:
5536:
5491:
5438:
5283:
5155:
5111:
4991:
4984:
4932:
4918:
3250:"A new Silurian fish close to the common ancestor of modern gnathostomes"
3090:
3072:
2393:
2113:
1969:
1935:
1896:
1881:
1858:
1794:
1671:
1632:
1547:
1303:
1275:
1171:
1090:
1061:
944:
793:
788:
778:
748:
520:
497:
351:
297:
37:
4278:
4233:
4097:
3981:
3714:"The oldest complete jawed vertebrates from the early Silurian of China"
3193:
1908:
1850:
1766:
1507:. When it was first discovered in 1980, it was originally regarded as a
1281:
Placoderms also share certain anatomical features only with the jawless
5898:
5797:
5642:
5579:
5524:
5506:
5305:
5296:
5082:
5053:
4998:
4960:
4904:
4876:
4455:
3917:
3080:
2933:
2896:
2801:
2426:
2187:
1863:
1786:
1660:
1430:
1396:
1384:
1355:
1348:
1201:
1159:
1079:
1028:
952:
925:
915:
904:
854:
833:
764:
738:
727:
711:
680:
603:
581:
577:
550:
531:
493:
469:
398:
237:
228:
184:
112:
82:
47:
3465:
3448:
1827:
5852:
5609:
5573:
5478:
5030:
4975:
4946:
4639:
4581:
4554:
3578:"The first vertebrate sexual organs evolved as an extra pair of legs"
3534:
2750:
2443:
2067:
1929:
1914:
1869:
1816:
1813:
1802:
1776:
1639:
1595:
1459:
1426:
1298:
1252:
1213:
1186:
1182:
1117:
1048:
1024:
982:
956:
932:
746:
of Western Australia, had streamlined, bullet-shaped head armor, and
658:
646:
588:
573:
538:
481:
477:
420:
260:
231:
154:
87:
31:
6005:
5194:
3868:
704:
6028:
5815:
5760:
5717:
5512:
5311:
5289:
5251:
5141:
5012:
5005:
4792:
4632:
4449:
4154:"New phyllolepids from Victoria and the relationships of the group"
2294:
1892:
1888:
1857:
was a thin fish that, when alive, looked vaguely like an elongated
1727:
1711:
1691:
1687:
1656:
1651:
1647:
1604:
1587:
1562:
1468:, the limbs were thick and short, while in advanced forms, such as
1438:
1363:
1312:
1209:
1205:
1194:
1121:
1020:
874:
773:
760:
672:
509:
505:
473:
431:
427:
313:
174:
103:
77:
72:
57:
52:
42:
3546:
1542:
1482:
1144:
The first officially described Silurian placoderm is an antiarch,
568:
of the early Silurian. They eventually outcompeted the previously
5903:
5443:
5433:
5415:
5104:
3171:
3152:
Colbert, Edwin H. (Edwin Harris); Knight, Charles Robert (1951).
2848:
1616:
1608:
1455:
1128:
1073:
843:
698:
584:
462:
412:
281:
127:
92:
67:
6063:
3826:
Recent Advances in the Origin and Early Radiation of Vertebrates
1038:
967:
5266:
4443:
4358:
Introduction to the Placodermi Extinct armored fishes with jaws
3620:
Pan, Zhaohui; Niu, Zhibin; Xian, Zumin; Zhu, Min (2023-01-03).
1924:
1891:
radiation. Critics of Janvier's position say that aside from a
1237:
1099:
1031:, as well as other, more primitive, groups. Earlier fossils of
684:
450:
423:
401:
164:
939:
5722:
4075:
3954:
Giles, Sam; Friedman, Matt; Brazeau, Martin D. (2015-01-12).
3375:
1917:
shows the interrelationships of placoderms according to Carr
1643:
1264:
1256:
1151:
1000:
489:
485:
1175:, also known from distinctively ornamented plates. In 2022,
591:), producing some of the first and most infamous vertebrate
5236:
1706:
1150:, which is known from distinctively ornamented plates from
446:
3604:
3522:
3348:
3155:
The dinosaur book: the ruling reptiles and their relatives
1899:, the two groups have little else in common anatomically.
1154:, China. It was originally considered to be from the late
4211:
3824:. In Arratia, G.; Wilson, M. V. H.; Cloutier, R. (eds.).
3540:
1297:. There are two further controversial orders: One is the
1260:
3487:
1293:
Currently, Placodermi are divided into eight recognized
3127:, then modern jawed vertebrates represent extant forms.
2708:
than others. The updated cladogram (Zhu et al., 2016):
1876:, as well as other paleontologists, has suggested that
1725:
and were found throughout the world. The petalichthids
763:, strongly supporting the idea that many, if not most,
4353:
Annetta Markussen-Brown, "Devonian Armoured Fish" 2000
4256:
3552:
1193:
appearance and diversity at the very beginning of the
1023:. At that time, they were already differentiated into
512:. 380-million-year-old fossils of three other genera,
4172:
Baltimore: The Johns Hopkins University Press, 1996.
4158:
Proceedings of the Linnean Society of New South Wales
3850:"Fish 380 Million Years Old Found With Unborn Embryo"
1228:
The earliest studies of placoderms were published by
1102:, China, known from an almost complete thoracic armor
5885:
5840:
5807:
5789:
5639:
5561:
5488:
5455:
5343:
5332:
5323:
5316:
5301:
5279:
5262:
3953:
3048:
3899:
3887:"Philippe Janvier Tree of Life Contributor Profile"
3797:
4422:
4170:The Rise of Fishes: 500 Million Years of Evolution
3376:Sallan, Lauren & Coates, Michael (June 2010).
4124:Oxford, New York: Oxford University Press, 1998.
484:), as well as the first vertebrates to have true
6117:
4020:
3940:
3828:. Munchen, Germany: Verlag Dr. Friedrich Pfeil.
3558:
560:The first identifiable placoderms appear in the
3819:"Placoderm anatomy and phylogeny: new insights"
3428:
3382:Proceedings of the National Academy of Sciences
1868:between its tubercles and the tubercles of the
1503:shield") was a long-snouted placoderm from the
1158:, although later study reconsidered its age at
457:(hence the name), and the rest of the body was
4182:
3528:
3344:
3342:
3304:
1166:plates are very similar to the early Devonian
5210:
4408:
4301:
4185:Annual Review of Earth and Planetary Sciences
3768:
3619:
3167:
3165:
3151:
1450:, particularly with later forms. The pair of
1446:sometimes naked rear portions often becoming
4476:
3668:
3816:
3711:
3369:
3339:
3247:
1619:and possibly claspers on the head as well.
894:
5217:
5203:
4415:
4401:
3162:
1518:, anatomical similarities in the brain of
767:were active swimmers, rather than passive
111:
4327:
3997:
3671:"The Antiarchi from Early Silurian Hunan"
3645:
3505:
3464:
3411:
3401:
3322:
3273:
1462:-like limbs. In primitive forms, such as
1288:
1035:placoderms have not yet been discovered.
530:, represent the oldest known examples of
3446:
1941:
1923:
1826:
1749:
1705:
1670:
1626:
1572:
1541:
1481:
1413:
1342:
1330:
1301:Stensioellida, containing the enigmatic
1181:is described from complete fossils from
1037:
966:
938:
914:
898:
703:
4014:
1745:
1694:or small, scaly arthrodires with blunt
1382:. Early arthrodires, such as the genus
6118:
1789:. As such, placoderm experts consider
1534:have also been inserted in the order.
719:One of the largest known arthrodires,
6010:
6009:
5224:
5198:
4436:
4396:
3893:
3867:Carr, Robert K.; et al. (2010).
3817:Goujet, Daniel; Young, Gavin (2004).
3662:
3221:
3219:
3158:. New York: McGraw-Hill. p. 153.
2704:, with some being more closer to the
1880:is not a placoderm, but instead is a
1400:measured just 15 cm. Fossils of
1307:; the other is the equally enigmatic
675:, many of which lived at or near the
617:358.9 million years ago, leaving the
4151:
4134:
3866:
3774:
3564:
3553:Long, Trinajstic & Johanson 2009
3434:
4205:10.1146/annurev-earth-040809-152416
4068:
3447:Engelman, Russell K. (2023-02-21).
1895:superficially similar to primitive
1477:
1321:
1223:
13:
3494:Frontiers in Ecology and Evolution
3351:Journal of Vertebrate Paleontology
3227:"Fossil reveals oldest live birth"
3216:
781:, such as the gigantic arthrodire
709:External anatomy of the placoderm
645:Many placoderms, particularly the
640:
14:
6172:
4346:
1701:
1686:("spine chests") were a group of
1666:
1433:. The order's name was coined by
1249:Swedish Museum of Natural History
931:, was a giant armoured placoderm
756:like that of other fast-swimming
541:, consisting of several distinct
5401:
5395:
5390:
5385:
5379:
5373:
5367:
5360:
5180:
5179:
4694:
3947:
3777:"Introduction to the Placodermi"
3065:
3051:
1822:
1568:
1014:
488:. They were also the first fish
468:Placoderms were among the first
141:
35:
3879:
3860:
3842:
3810:
3791:
3705:
3613:
3595:
3570:
3481:
3440:
1615:, with the males having pelvic
1537:
861:The placoderm claspers are not
697:and were made of both bone and
3363:10.1080/02724634.1999.10011135
3298:
3241:
3145:
3113:
1793:to be the sister group of the
1756:Pseudopetalicthys problematica
1065:(and then, the discoveries of
887:showcases that the vertebrate
1:
6156:Taxa named by Frederick McCoy
3580:. 8 June 2014. Archived from
3134:
1739:Quasipetalichthys haikouensis
1578:Kimbryanodus williamburyensis
1454:were modified into a pair of
1326:
537:Placoderms are thought to be
6146:Llandovery first appearances
5962:List of transitional fossils
3139:
1902:
1622:
1409:
903:Evolution and extinction of
453:were covered by articulated
7:
3123:in relation to the rest of
3044:
629:survivors who subsequently
10:
6177:
5242:Timeline of fish evolution
3738:10.1038/s41586-022-05136-8
3529:Long & Trinajstic 2010
1906:
988:
907:. The diagram is based on
891:evolved from gill arches.
461:or naked depending on the
6018:
5996:
5970:
5930:Lists of prehistoric fish
5922:
5831:
5780:
5771:
5679:
5638:
5560:
5551:
5487:
5454:
5424:
5414:
5358:
5250:
5232:
5176:
5075:
5052:
5029:
4974:
4850:
4841:
4768:
4734:
4712:
4703:
4692:
4663:
4577:
4562:
4550:
4546:
4527:
4503:
4485:
4472:
4432:
3626:Earth System Science Data
3275:10.1016/j.cub.2021.05.053
2947:
2927:
2920:
2910:
2893:
2886:
2869:
2862:
2845:
2838:
2814:
2799:
2792:
2785:
2770:
2763:
2748:
2741:
2723:
2716:
2574:
2557:
2540:
2523:
2516:
2499:
2492:
2475:
2440:
2423:
2416:
2390:
2373:
2366:
2359:
2352:
2325:
2308:
2291:
2284:
2277:
2259:
2252:
2237:
2230:
2215:
2198:
2191:
2164:
2149:
2142:
2135:
2111:
2094:
2087:
2080:
2065:
2058:
2034:
2017:
2010:
1993:
1986:
1966:
1959:
1441:, mistakenly thought the
924:, among the first of the
825:Materpiscis attenboroughi
367:
360:
225:
220:
138:Scientific classification
136:
119:
110:
23:
6131:Prehistoric fish classes
4135:Long, J. A. (May 1983).
3507:10.3389/fevo.2022.969158
3106:
2932:(total group, including
1521:Brindabellaspis stensioi
1488:Brindabellaspis stensioi
1009:end-Devonian extinctions
895:Evolution and extinction
5988:Vertebrate paleontology
4364:"PALAEOZOIC FOSSILS UK"
4043:10.1126/science.aah3764
3647:10.5194/essd-15-41-2023
3403:10.1073/pnas.0914000107
1646:fish") were flattened,
1420:Bothriolepis canadensis
1243:In the late 1920s, Dr.
480:from the first pair of
276:"Maxillate placoderms"
16:Extinct Class of Fishes
5083:"maxillate placoderms"
4320:10.1098/rsbl.2011.1033
3678:Vertebrata PalAsiatica
1951:
1939:
1836:
1759:
1715:
1680:
1636:
1580:
1551:
1491:
1423:
1352:
1340:
1289:Taxonomy and phylogeny
1132:, a close relative of
1056:
986:
964:
936:
912:
716:
679:. Many, primarily the
6098:Paleobiology Database
4387:Dunkleosteus terrelli
3906:Journal of Morphology
3902:Cowralepis mclachlani
3800:Journal of Morphology
3669:Wang Junqing (1991).
3324:10.1093/sysbio/syw107
1945:
1927:
1909:Agnatha § Groups
1830:
1753:
1709:
1674:
1630:
1576:
1545:
1485:
1417:
1346:
1334:
1185:, late Llandovery of
1041:
970:
942:
918:
902:
865:with the claspers in
787:, various members of
707:
6151:Devonian extinctions
5983:Transitional fossils
5622:Pseudopetalichthyida
4770:Macropetalichthyidae
4736:Quasipetalichthyidae
4718:Diandongpetalichthys
4380:Placodermi: Overview
3260:(16): 3613–3620.e2.
1948:Diandongpetalichthys
1791:Pseudopetalichthyida
1763:Pseudopetalichthyida
1746:Pseudopetalichthyida
1655:placoderms, such as
1309:Pseudopetalichthyida
871:convergent evolution
867:cartilaginous fishes
839:Incisoscutum ritchei
712:Coccosteus decipiens
549:to all living jawed
496:, the second set of
345:Pseudopetalichthyida
123:Bothriolepis panderi
6161:Paraphyletic groups
4370:on 17 October 2012.
4279:10.1038/nature07732
4271:2009Natur.457.1124L
4265:(7233): 1124–1127.
4234:10.1038/nature06966
4226:2008Natur.453..650L
4197:2010AREPS..38..255L
4152:Long, J.A. (1984).
4098:10.1038/nature08176
4090:2009Natur.460..888A
4035:2016Sci...354..334Z
3982:10.1038/nature14065
3974:2015Natur.520...82G
3730:2022Natur.609..954Z
3638:2023ESSD...15...41P
3394:2010PNAS..10710131S
3388:(22): 10131–10135.
3266:2021CBio...31E3613L
3229:. BBC. May 28, 2008
3194:10.1038/nature11555
3186:2012Natur.491..748R
3059:Paleontology portal
1853:slates of Germany.
1833:Stensioella heintzi
1782:Stensioella heintzi
1772:Stensioella heintzi
1524:and the brain of a
1435:Edward Drinker Cope
1240:-like vertebrates.
1136:that further links
1088:the basal antiarch
5671:Ischnacanthiformes
5519:Cephalaspidomorphi
5149:Pseudopetalichthys
4665:Weejasperaspididae
4579:Palaeacanthaspidae
3918:10.1002/jmor.10719
3311:Systematic Biology
3086:List of placoderms
2936:as stem lineages)
2726:Cephalaspidomorphi
1997:Pseudopetalichthys
1952:
1940:
1837:
1760:
1716:
1681:
1650:, bottom-dwelling
1637:
1613:sexually dimorphic
1581:
1552:
1548:Phyllolepis orvini
1492:
1424:
1353:
1341:
1057:
987:
965:
937:
913:
830:David Attenborough
717:
381:(from Greek πλάξ (
6113:
6112:
6085:Open Tree of Life
6012:Taxon identifiers
6003:
6002:
5918:
5917:
5914:
5913:
5845:Palaeonisciformes
5793:Onychodontiformes
5750:Petalodontiformes
5744:Iniopterygiformes
5707:Phoebodontiformes
5695:Ctenacanthiformes
5665:Diplacanthiformes
5547:
5546:
5320:Myllokunmingiidae
5226:Evolution of fish
5192:
5191:
5172:
5171:
5122:
5121:
4970:
4969:
4837:
4836:
4800:Macropetalichthys
4759:Quasipetalichthys
4752:Pauropetalichthys
4690:
4689:
4542:
4541:
4538:
4537:
4523:
4522:
4220:(7195): 650–652.
4122:Early Vertebrates
4084:(7257): 888–889.
4029:(6310): 334–336.
3724:(7929): 954–958.
3466:10.3390/d15030318
3180:(7426): 748–751.
3041:
3040:
3032:
3031:
3023:
3022:
3014:
3013:
3005:
3004:
2996:
2995:
2987:
2986:
2978:
2977:
2969:
2968:
2960:
2959:
2827:
2826:
2697:
2696:
2688:
2687:
2679:
2678:
2670:
2669:
2661:
2660:
2652:
2651:
2643:
2642:
2634:
2633:
2625:
2624:
2616:
2615:
2607:
2606:
2598:
2597:
2589:
2588:
2464:
2463:
2455:
2454:
2405:
2404:
2338:
2337:
2177:
2176:
2124:
2123:
2047:
2046:
991:Evolution of fish
849:Austrophyllolepis
828:(after scientist
376:
375:
216:
6168:
6106:
6105:
6093:
6092:
6080:
6079:
6067:
6066:
6054:
6053:
6052:
6039:
6038:
6037:
6007:
6006:
5978:Prehistoric life
5889:Pycnodontiformes
5887:
5872:Acipenseriformes
5842:
5809:
5791:
5778:
5777:
5641:
5586:Brindabellaspida
5563:
5558:
5557:
5501:Pteraspidomorphi
5490:
5457:
5422:
5421:
5406:
5405:
5399:
5394:
5389:
5384:
5383:
5377:
5372:
5371:
5364:
5345:
5334:
5325:
5318:
5303:
5281:
5264:
5219:
5212:
5205:
5196:
5195:
5183:
5182:
5135:Holopetalichthys
5091:Bianchengichthys
5080:
5079:
4848:
4847:
4807:Notopetalichthys
4779:Ellopetalichthys
4710:
4709:
4698:
4626:Palaeacanthaspis
4560:
4559:
4548:
4547:
4483:
4482:
4474:
4473:
4467:
4466:
4434:
4433:
4417:
4410:
4403:
4394:
4393:
4371:
4366:. Archived from
4341:
4331:
4298:
4253:
4208:
4165:
4148:
4117:
4069:Other references
4063:
4062:
4018:
4012:
4011:
4001:
3952:
3951:
3944:
3938:
3937:
3897:
3891:
3890:
3883:
3877:
3876:
3864:
3858:
3857:
3846:
3840:
3839:
3823:
3814:
3808:
3807:
3795:
3789:
3788:
3786:
3784:
3772:
3766:
3765:
3709:
3703:
3702:
3700:
3694:. Archived from
3690:
3675:
3666:
3660:
3659:
3649:
3617:
3611:
3608:
3602:
3599:
3593:
3592:
3590:
3589:
3574:
3568:
3562:
3556:
3550:
3544:
3541:Long et al. 2008
3538:
3532:
3526:
3520:
3519:
3509:
3485:
3479:
3478:
3468:
3444:
3438:
3432:
3426:
3425:
3415:
3405:
3373:
3367:
3366:
3346:
3337:
3336:
3326:
3302:
3296:
3295:
3277:
3245:
3239:
3238:
3236:
3234:
3223:
3214:
3213:
3169:
3160:
3159:
3149:
3128:
3117:
3075:
3070:
3069:
3068:
3061:
3056:
3055:
3054:
2923:
2922:
2913:
2912:
2889:
2888:
2865:
2864:
2841:
2840:
2795:
2794:
2788:
2787:
2766:
2765:
2744:
2743:
2719:
2718:
2712:
2711:
2519:
2518:
2495:
2494:
2419:
2418:
2369:
2368:
2362:
2361:
2355:
2354:
2287:
2286:
2280:
2279:
2255:
2254:
2233:
2232:
2194:
2193:
2145:
2144:
2138:
2137:
2090:
2089:
2083:
2082:
2061:
2060:
2013:
2012:
1989:
1988:
1962:
1961:
1955:
1954:
1874:Philippe Janvier
1769:, Germany. Like
1677:Palaeacanthaspis
1592:acanthothoracids
1516:Philippe Janvier
1478:Brindabellaspida
1322:Placoderm orders
1224:History of study
957:Middle Famennian
821:
820:
816:
813:
802:Fitzroy Crossing
777:, and some were
615:Hangenberg event
607:and the massive
566:Llandovery epoch
564:during the late
417:prehistoric fish
306:Bianchengichthys
211:
206:
146:
145:
115:
97:
34:
27:Temporal range:
21:
20:
6176:
6175:
6171:
6170:
6169:
6167:
6166:
6165:
6116:
6115:
6114:
6109:
6101:
6096:
6088:
6083:
6075:
6070:
6062:
6057:
6048:
6047:
6042:
6033:
6032:
6027:
6014:
6004:
5999:
5992:
5966:
5910:
5858:Polypteriformes
5827:
5821:Tetrapodomorpha
5767:
5701:Hybodontiformes
5675:
5652:Acanthodiformes
5634:
5543:
5483:
5473:Paraconodontida
5450:
5418:
5410:
5409:
5408:
5400:
5378:
5366:
5356:
5275:Cephalochordate
5246:
5228:
5223:
5193:
5188:
5168:
5128:Brindabellaspis
5118:
5071:
5048:
5025:
4966:
4856:Austroptyctodus
4833:
4786:Epipetalichthys
4764:
4730:
4725:Neopetalichthys
4699:
4686:
4659:
4573:
4534:
4519:
4499:
4468:
4439:
4438:
4428:
4421:
4362:
4349:
4344:
4308:Biology Letters
4071:
4066:
4019:
4015:
3968:(7545): 82–85.
3946:
3945:
3941:
3898:
3894:
3885:
3884:
3880:
3865:
3861:
3856:. June 6, 2008.
3848:
3847:
3843:
3836:
3821:
3815:
3811:
3796:
3792:
3782:
3780:
3775:Waggoner, Ben.
3773:
3769:
3710:
3706:
3698:
3688:
3673:
3667:
3663:
3618:
3614:
3609:
3605:
3600:
3596:
3587:
3585:
3576:
3575:
3571:
3563:
3559:
3551:
3547:
3539:
3535:
3527:
3523:
3486:
3482:
3445:
3441:
3433:
3429:
3374:
3370:
3347:
3340:
3303:
3299:
3254:Current Biology
3246:
3242:
3232:
3230:
3225:
3224:
3217:
3170:
3163:
3150:
3146:
3142:
3137:
3132:
3131:
3118:
3114:
3109:
3071:
3066:
3064:
3057:
3052:
3050:
3047:
3042:
3033:
3024:
3015:
3006:
2997:
2988:
2979:
2970:
2961:
2917:Eugnathostomata
2828:
2706:Eugnathostomata
2698:
2689:
2680:
2671:
2662:
2653:
2644:
2635:
2626:
2617:
2608:
2599:
2590:
2465:
2456:
2406:
2349:Eubrachythoraci
2339:
2178:
2125:
2048:
2021:Brindabellaspis
1911:
1905:
1825:
1748:
1704:
1669:
1625:
1571:
1540:
1512:acanthothoracid
1496:Brindabellaspis
1480:
1412:
1368:bottom dwellers
1329:
1324:
1291:
1269:chondrichthyans
1226:
1017:
993:
979:Middle Devonian
897:
889:shoulder girdle
818:
814:
811:
809:
715:
683:, were active,
643:
641:Characteristics
527:Austroptyctodus
455:armoured plates
338:Brindabellaspis
321:Eugnathostomata
210:
204:
140:
106:
96:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
40:
30:439–358.9
29:
28:
25:
17:
12:
11:
5:
6174:
6164:
6163:
6158:
6153:
6148:
6143:
6138:
6133:
6128:
6111:
6110:
6108:
6107:
6094:
6081:
6068:
6055:
6040:
6024:
6022:
6016:
6015:
6001:
6000:
5997:
5994:
5993:
5991:
5990:
5985:
5980:
5974:
5972:
5968:
5967:
5965:
5964:
5959:
5958:
5957:
5952:
5947:
5942:
5937:
5926:
5924:
5920:
5919:
5916:
5915:
5912:
5911:
5909:
5908:
5907:
5906:
5901:
5896:
5891:
5877:
5876:
5875:
5874:
5863:
5862:
5861:
5860:
5849:
5848:
5837:
5835:
5829:
5828:
5826:
5825:
5824:
5823:
5818:
5813:
5811:Porolepiformes
5800:
5795:
5786:
5784:
5775:
5769:
5768:
5766:
5765:
5764:
5763:
5758:
5756:Symmoriiformes
5752:
5746:
5740:
5738:Eugeneodontida
5728:
5727:
5726:
5725:
5720:
5715:
5709:
5703:
5697:
5689:Elasmobranchii
5685:
5683:
5677:
5676:
5674:
5673:
5667:
5661:
5658:Climatiiformes
5654:
5647:
5645:
5636:
5635:
5633:
5632:
5625:
5618:
5616:Acanthothoraci
5612:
5606:
5600:
5594:
5592:Petalichthyida
5588:
5582:
5576:
5569:
5567:
5555:
5549:
5548:
5545:
5544:
5542:
5541:
5540:
5539:
5533:
5527:
5515:
5509:
5503:
5496:
5494:
5485:
5484:
5482:
5481:
5475:
5469:
5466:Protoconodonta
5461:
5459:
5452:
5451:
5449:
5448:
5447:
5446:
5436:
5430:
5428:
5419:
5412:
5411:
5359:
5357:
5355:
5354:
5353:
5352:
5341:
5337:Myllokunmingia
5314:
5309:
5294:
5293:
5292:
5287:
5272:
5271:
5270:
5256:
5254:
5248:
5247:
5245:
5244:
5239:
5233:
5230:
5229:
5222:
5221:
5214:
5207:
5199:
5190:
5189:
5187:
5177:
5174:
5173:
5170:
5169:
5167:
5166:
5159:
5152:
5145:
5138:
5131:
5123:
5120:
5119:
5117:
5116:
5108:
5101:
5094:
5086:
5084:
5077:
5073:
5072:
5070:
5069:
5068:
5067:
5058:
5056:
5050:
5049:
5047:
5046:
5045:
5044:
5035:
5033:
5027:
5026:
5024:
5023:
5016:
5009:
5002:
4995:
4988:
4980:
4978:
4972:
4971:
4968:
4967:
4965:
4964:
4957:
4950:
4943:
4936:
4929:
4922:
4915:
4908:
4901:
4894:
4887:
4880:
4873:
4866:
4859:
4851:
4845:
4839:
4838:
4835:
4834:
4832:
4831:
4824:
4817:
4810:
4803:
4796:
4789:
4782:
4774:
4772:
4766:
4765:
4763:
4762:
4755:
4748:
4740:
4738:
4732:
4731:
4729:
4728:
4721:
4713:
4707:
4705:Petalichthyida
4701:
4700:
4693:
4691:
4688:
4687:
4685:
4684:
4681:Weejasperaspis
4677:
4674:Murrindalaspis
4669:
4667:
4661:
4660:
4658:
4657:
4650:
4643:
4636:
4629:
4622:
4615:
4608:
4601:
4594:
4586:
4584:
4575:
4574:
4572:
4571:
4563:
4557:
4552:Acanthothoraci
4544:
4543:
4540:
4539:
4536:
4535:
4528:
4525:
4524:
4521:
4520:
4518:
4517:
4516:
4515:
4509:
4507:
4501:
4500:
4498:
4497:
4496:
4495:
4486:
4480:
4470:
4469:
4465:
4464:
4458:
4452:
4446:
4430:
4429:
4420:
4419:
4412:
4405:
4397:
4391:
4390:
4385:BBC—report on
4382:
4377:
4372:
4360:
4355:
4348:
4347:External links
4345:
4343:
4342:
4314:(3): 453–456.
4299:
4254:
4209:
4180:
4166:
4149:
4132:
4118:
4072:
4070:
4067:
4065:
4064:
4013:
3939:
3912:(7): 775–804.
3892:
3878:
3859:
3841:
3834:
3809:
3790:
3767:
3704:
3701:on 2013-12-12.
3684:(3): 240–244.
3661:
3612:
3603:
3594:
3569:
3557:
3545:
3533:
3521:
3480:
3439:
3427:
3368:
3357:(2): 204–219.
3338:
3317:(4): 499–516.
3297:
3240:
3215:
3161:
3143:
3141:
3138:
3136:
3133:
3130:
3129:
3111:
3110:
3108:
3105:
3104:
3103:
3098:
3096:Chondrichthyes
3093:
3088:
3083:
3077:
3076:
3062:
3046:
3043:
3039:
3038:
3035:
3034:
3030:
3029:
3026:
3025:
3021:
3020:
3017:
3016:
3012:
3011:
3008:
3007:
3003:
3002:
2999:
2998:
2994:
2993:
2990:
2989:
2985:
2984:
2981:
2980:
2976:
2975:
2972:
2971:
2967:
2966:
2963:
2962:
2958:
2957:
2954:
2953:
2946:
2943:
2942:
2939:
2938:
2930:Chondrichthyes
2926:
2921:
2919:
2911:
2909:
2906:
2905:
2902:
2901:
2892:
2887:
2885:
2882:
2881:
2878:
2877:
2868:
2863:
2861:
2858:
2857:
2854:
2853:
2844:
2839:
2837:
2834:
2833:
2830:
2829:
2825:
2824:
2821:
2820:
2813:
2810:
2809:
2806:
2805:
2798:
2793:
2791:
2786:
2784:
2781:
2780:
2777:
2776:
2773:Petalichthyida
2769:
2764:
2762:
2759:
2758:
2755:
2754:
2747:
2742:
2740:
2734:
2733:
2730:
2729:
2722:
2717:
2715:
2710:
2695:
2694:
2691:
2690:
2686:
2685:
2682:
2681:
2677:
2676:
2673:
2672:
2668:
2667:
2664:
2663:
2659:
2658:
2655:
2654:
2650:
2649:
2646:
2645:
2641:
2640:
2637:
2636:
2632:
2631:
2628:
2627:
2623:
2622:
2619:
2618:
2614:
2613:
2610:
2609:
2605:
2604:
2601:
2600:
2596:
2595:
2592:
2591:
2587:
2586:
2583:
2582:
2573:
2570:
2569:
2566:
2565:
2556:
2553:
2552:
2549:
2548:
2539:
2536:
2535:
2532:
2531:
2522:
2517:
2515:
2512:
2511:
2508:
2507:
2498:
2493:
2491:
2488:
2487:
2484:
2483:
2474:
2471:
2470:
2467:
2466:
2462:
2461:
2458:
2457:
2453:
2452:
2449:
2448:
2439:
2436:
2435:
2432:
2431:
2422:
2417:
2415:
2412:
2411:
2408:
2407:
2403:
2402:
2399:
2398:
2389:
2386:
2385:
2382:
2381:
2372:
2367:
2365:
2360:
2358:
2353:
2351:
2345:
2344:
2341:
2340:
2336:
2335:
2332:
2331:
2328:Buchanosteidae
2324:
2321:
2320:
2317:
2316:
2307:
2304:
2303:
2300:
2299:
2290:
2285:
2283:
2278:
2276:
2270:
2269:
2266:
2265:
2258:
2253:
2251:
2248:
2247:
2244:
2243:
2236:
2231:
2229:
2226:
2225:
2222:
2221:
2214:
2211:
2210:
2207:
2206:
2202:Wuttagoonaspis
2197:
2192:
2190:
2184:
2183:
2180:
2179:
2175:
2174:
2171:
2170:
2163:
2160:
2159:
2156:
2155:
2152:Petalichthyida
2148:
2143:
2141:
2136:
2134:
2131:
2130:
2127:
2126:
2122:
2121:
2118:
2117:
2110:
2107:
2106:
2103:
2102:
2093:
2088:
2086:
2081:
2079:
2076:
2075:
2072:
2071:
2064:
2059:
2057:
2054:
2053:
2050:
2049:
2045:
2044:
2041:
2040:
2037:Acanthothoraci
2033:
2030:
2029:
2026:
2025:
2016:
2011:
2009:
2006:
2005:
2002:
2001:
1992:
1987:
1985:
1979:
1978:
1975:
1974:
1965:
1960:
1958:
1953:
1913:The following
1904:
1901:
1897:holocephalians
1886:Chondrichthyan
1861:, or a skinny
1848:Lower Devonian
1824:
1821:
1810:Acanthothoraci
1747:
1744:
1723:Early Devonian
1719:Petalichthyida
1703:
1702:Petalichthyida
1700:
1684:Acanthothoraci
1668:
1667:Acanthothoraci
1665:
1624:
1621:
1600:holocephalians
1570:
1567:
1539:
1536:
1531:incertae sedis
1509:weejasperaspid
1505:Early Devonian
1479:
1476:
1411:
1408:
1360:apex predators
1328:
1325:
1323:
1320:
1290:
1287:
1225:
1222:
1142:
1141:
1125:
1113:incertae sedis
1110:, a placoderm
1103:
1053:Early Silurian
1016:
1013:
929:apex predators
909:Michael Benton
896:
893:
769:ambush-hunters
744:Gogo Formation
742:, both of the
708:
651:Petalichthyida
642:
639:
627:chondrichthyan
593:apex predators
437:. While their
397:) 'skin') are
374:
373:
372:
371:
365:
364:
358:
357:
356:
355:
347:
341:
333:
325:
324:
323:
317:
309:
301:
293:
285:
274:
267:Acanthothoraci
263:
257:
251:
249:Petalichthyida
245:
234:
223:
222:
218:
217:
202:
198:
197:
192:
188:
187:
182:
178:
177:
172:
168:
167:
162:
158:
157:
152:
148:
147:
134:
133:
117:
116:
108:
107:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
41:
36:
26:
15:
9:
6:
4:
3:
2:
6173:
6162:
6159:
6157:
6154:
6152:
6149:
6147:
6144:
6142:
6141:Silurian fish
6139:
6137:
6136:Devonian fish
6134:
6132:
6129:
6127:
6124:
6123:
6121:
6104:
6099:
6095:
6091:
6086:
6082:
6078:
6073:
6069:
6065:
6060:
6056:
6051:
6045:
6041:
6036:
6030:
6026:
6025:
6023:
6021:
6017:
6013:
6008:
5995:
5989:
5986:
5984:
5981:
5979:
5976:
5975:
5973:
5969:
5963:
5960:
5956:
5953:
5951:
5948:
5946:
5945:cartilaginous
5943:
5941:
5938:
5936:
5933:
5932:
5931:
5928:
5927:
5925:
5921:
5905:
5902:
5900:
5897:
5895:
5892:
5890:
5884:
5883:
5882:
5879:
5878:
5873:
5870:
5869:
5868:
5865:
5864:
5859:
5856:
5855:
5854:
5851:
5850:
5846:
5839:
5838:
5836:
5834:
5830:
5822:
5819:
5817:
5814:
5812:
5806:
5805:
5804:
5801:
5799:
5796:
5794:
5788:
5787:
5785:
5783:
5779:
5776:
5774:
5770:
5762:
5759:
5757:
5753:
5751:
5747:
5745:
5741:
5739:
5735:
5734:
5733:
5730:
5729:
5724:
5721:
5719:
5716:
5714:
5710:
5708:
5704:
5702:
5698:
5696:
5692:
5691:
5690:
5687:
5686:
5684:
5682:
5681:Cartilaginous
5678:
5672:
5668:
5666:
5662:
5659:
5655:
5653:
5649:
5648:
5646:
5644:
5637:
5630:
5629:Stensioellida
5626:
5623:
5619:
5617:
5613:
5611:
5607:
5605:
5604:Ptyctodontida
5601:
5599:
5595:
5593:
5589:
5587:
5583:
5581:
5577:
5575:
5571:
5570:
5568:
5566:
5559:
5556:
5554:
5550:
5538:
5534:
5532:
5528:
5526:
5522:
5521:
5520:
5516:
5514:
5510:
5508:
5504:
5502:
5498:
5497:
5495:
5493:
5486:
5480:
5476:
5474:
5470:
5467:
5463:
5462:
5460:
5453:
5445:
5442:
5441:
5440:
5437:
5435:
5432:
5431:
5429:
5427:
5423:
5420:
5417:
5413:
5407:
5404:
5398:
5393:
5388:
5382:
5376:
5370:
5363:
5350:
5349:
5348:Zhongxiniscus
5342:
5339:
5338:
5330:
5329:
5328:Haikouichthys
5321:
5315:
5313:
5310:
5308:
5307:
5300:
5299:
5298:
5295:
5291:
5288:
5286:
5285:
5278:
5277:
5276:
5273:
5269:
5268:
5261:
5260:
5259:Basal member
5258:
5257:
5255:
5253:
5249:
5243:
5240:
5238:
5235:
5234:
5231:
5227:
5220:
5215:
5213:
5208:
5206:
5201:
5200:
5197:
5186:
5178:
5175:
5165:
5164:
5163:Xiushanosteus
5160:
5158:
5157:
5153:
5151:
5150:
5146:
5144:
5143:
5139:
5137:
5136:
5132:
5130:
5129:
5125:
5124:
5114:
5113:
5109:
5107:
5106:
5102:
5100:
5099:
5098:Entelognathus
5095:
5093:
5092:
5088:
5087:
5085:
5081:
5078:
5074:
5066:
5062:
5061:
5060:
5059:
5057:
5055:
5051:
5043:
5039:
5038:
5037:
5036:
5034:
5032:
5028:
5022:
5021:
5017:
5015:
5014:
5010:
5008:
5007:
5003:
5001:
5000:
4996:
4994:
4993:
4989:
4987:
4986:
4982:
4981:
4979:
4977:
4973:
4963:
4962:
4958:
4956:
4955:
4954:Rhamphodopsis
4951:
4949:
4948:
4944:
4942:
4941:
4937:
4935:
4934:
4930:
4928:
4927:
4923:
4921:
4920:
4916:
4914:
4913:
4909:
4907:
4906:
4902:
4900:
4899:
4898:Eczematolepis
4895:
4893:
4892:
4891:Destnoporella
4888:
4886:
4885:
4881:
4879:
4878:
4874:
4872:
4871:
4867:
4865:
4864:
4860:
4858:
4857:
4853:
4852:
4849:
4846:
4844:
4843:Ptyctodontida
4840:
4830:
4829:
4825:
4823:
4822:
4821:Tongdzuylepis
4818:
4816:
4815:
4814:Shearsbyaspis
4811:
4809:
4808:
4804:
4802:
4801:
4797:
4795:
4794:
4790:
4788:
4787:
4783:
4781:
4780:
4776:
4775:
4773:
4771:
4767:
4761:
4760:
4756:
4754:
4753:
4749:
4747:
4746:
4742:
4741:
4739:
4737:
4733:
4727:
4726:
4722:
4720:
4719:
4715:
4714:
4711:
4708:
4706:
4702:
4697:
4683:
4682:
4678:
4676:
4675:
4671:
4670:
4668:
4666:
4662:
4656:
4655:
4651:
4649:
4648:
4644:
4642:
4641:
4637:
4635:
4634:
4630:
4628:
4627:
4623:
4621:
4620:
4616:
4614:
4613:
4609:
4607:
4606:
4602:
4600:
4599:
4595:
4593:
4592:
4588:
4587:
4585:
4583:
4580:
4576:
4570:
4569:
4565:
4564:
4561:
4558:
4556:
4553:
4549:
4545:
4533:
4532:
4526:
4513:
4512:
4511:
4510:
4508:
4506:
4502:
4494:
4493:Gnathostomata
4490:
4489:
4488:
4487:
4484:
4481:
4479:
4478:Gnathostomata
4475:
4471:
4463:
4462:Gnathostomata
4460:Infraphylum:
4459:
4457:
4453:
4451:
4447:
4445:
4441:
4440:
4435:
4431:
4426:
4418:
4413:
4411:
4406:
4404:
4399:
4398:
4395:
4389:
4388:
4383:
4381:
4378:
4376:
4373:
4369:
4365:
4361:
4359:
4356:
4354:
4351:
4350:
4339:
4335:
4330:
4325:
4321:
4317:
4313:
4309:
4305:
4300:
4296:
4292:
4288:
4284:
4280:
4276:
4272:
4268:
4264:
4260:
4255:
4251:
4247:
4243:
4239:
4235:
4231:
4227:
4223:
4219:
4215:
4210:
4206:
4202:
4198:
4194:
4190:
4186:
4181:
4179:
4178:0-8018-5438-5
4175:
4171:
4167:
4163:
4159:
4155:
4150:
4147:(2): 295–320.
4146:
4142:
4141:Palaeontology
4138:
4133:
4131:
4130:0-19-854047-7
4127:
4123:
4119:
4115:
4111:
4107:
4103:
4099:
4095:
4091:
4087:
4083:
4079:
4074:
4073:
4060:
4056:
4052:
4048:
4044:
4040:
4036:
4032:
4028:
4024:
4017:
4009:
4005:
4000:
3995:
3991:
3987:
3983:
3979:
3975:
3971:
3967:
3963:
3962:
3957:
3950:
3943:
3935:
3931:
3927:
3923:
3919:
3915:
3911:
3907:
3903:
3896:
3888:
3882:
3874:
3870:
3863:
3855:
3854:Science Daily
3851:
3845:
3837:
3835:3-89937-052-X
3831:
3827:
3820:
3813:
3805:
3801:
3794:
3778:
3771:
3763:
3759:
3755:
3751:
3747:
3743:
3739:
3735:
3731:
3727:
3723:
3719:
3715:
3708:
3697:
3693:
3687:
3683:
3679:
3672:
3665:
3657:
3653:
3648:
3643:
3639:
3635:
3631:
3627:
3623:
3616:
3607:
3598:
3584:on 2016-12-20
3583:
3579:
3573:
3566:
3561:
3554:
3549:
3542:
3537:
3530:
3525:
3517:
3513:
3508:
3503:
3499:
3495:
3491:
3484:
3476:
3472:
3467:
3462:
3458:
3454:
3450:
3443:
3436:
3431:
3423:
3419:
3414:
3409:
3404:
3399:
3395:
3391:
3387:
3383:
3379:
3372:
3364:
3360:
3356:
3352:
3345:
3343:
3334:
3330:
3325:
3320:
3316:
3312:
3308:
3301:
3293:
3289:
3285:
3281:
3276:
3271:
3267:
3263:
3259:
3255:
3251:
3244:
3228:
3222:
3220:
3211:
3207:
3203:
3199:
3195:
3191:
3187:
3183:
3179:
3175:
3168:
3166:
3157:
3156:
3148:
3144:
3126:
3125:Gnathostomata
3122:
3116:
3112:
3102:
3101:Entelognathus
3099:
3097:
3094:
3092:
3089:
3087:
3084:
3082:
3079:
3078:
3074:
3063:
3060:
3049:
3037:
3036:
3028:
3027:
3019:
3018:
3010:
3009:
3001:
3000:
2992:
2991:
2983:
2982:
2974:
2973:
2965:
2964:
2956:
2955:
2952:
2951:
2945:
2944:
2941:
2940:
2937:
2935:
2931:
2925:
2924:
2918:
2915:
2914:
2908:
2907:
2904:
2903:
2900:
2899:
2898:
2891:
2890:
2884:
2883:
2880:
2879:
2876:
2875:
2874:
2873:Entelognathus
2867:
2866:
2860:
2859:
2856:
2855:
2852:
2851:
2850:
2843:
2842:
2836:
2835:
2832:
2831:
2823:
2822:
2819:
2818:
2817:Ptyctodontida
2812:
2811:
2808:
2807:
2804:
2803:
2797:
2796:
2790:
2789:
2783:
2782:
2779:
2778:
2775:
2774:
2768:
2767:
2761:
2760:
2757:
2756:
2753:
2752:
2746:
2745:
2739:
2738:Gnathostomata
2736:
2735:
2732:
2731:
2728:
2727:
2721:
2720:
2714:
2713:
2709:
2707:
2703:
2693:
2692:
2684:
2683:
2675:
2674:
2666:
2665:
2657:
2656:
2648:
2647:
2639:
2638:
2630:
2629:
2621:
2620:
2612:
2611:
2603:
2602:
2594:
2593:
2585:
2584:
2581:
2580:
2579:
2572:
2571:
2568:
2567:
2564:
2563:
2562:
2561:Gorgonichthys
2555:
2554:
2551:
2550:
2547:
2546:
2545:
2544:Erromenosteus
2538:
2537:
2534:
2533:
2530:
2529:
2528:
2521:
2520:
2514:
2513:
2510:
2509:
2506:
2505:
2504:
2497:
2496:
2490:
2489:
2486:
2485:
2482:
2481:
2480:
2473:
2472:
2469:
2468:
2460:
2459:
2451:
2450:
2447:
2446:
2445:
2438:
2437:
2434:
2433:
2430:
2429:
2428:
2421:
2420:
2414:
2413:
2410:
2409:
2401:
2400:
2397:
2396:
2395:
2388:
2387:
2384:
2383:
2380:
2379:
2378:
2371:
2370:
2364:
2363:
2357:
2356:
2350:
2347:
2346:
2343:
2342:
2334:
2333:
2330:
2329:
2323:
2322:
2319:
2318:
2315:
2314:
2313:
2306:
2305:
2302:
2301:
2298:
2297:
2296:
2289:
2288:
2282:
2281:
2275:
2274:Brachythoraci
2272:
2271:
2268:
2267:
2264:
2263:
2262:Phlyctaeniida
2257:
2256:
2250:
2249:
2246:
2245:
2242:
2241:
2235:
2234:
2228:
2227:
2224:
2223:
2220:
2219:
2218:Actinolepidae
2213:
2212:
2209:
2208:
2205:
2204:
2203:
2196:
2195:
2189:
2186:
2185:
2182:
2181:
2173:
2172:
2169:
2168:
2167:Ptyctodontida
2162:
2161:
2158:
2157:
2154:
2153:
2147:
2146:
2140:
2139:
2133:
2132:
2129:
2128:
2120:
2119:
2116:
2115:
2109:
2108:
2105:
2104:
2101:
2100:
2099:
2092:
2091:
2085:
2084:
2078:
2077:
2074:
2073:
2070:
2069:
2063:
2062:
2056:
2055:
2052:
2051:
2043:
2042:
2039:
2038:
2032:
2031:
2028:
2027:
2024:
2023:
2022:
2015:
2014:
2008:
2007:
2004:
2003:
2000:
1999:
1998:
1991:
1990:
1984:
1981:
1980:
1977:
1976:
1973:
1972:
1971:
1964:
1963:
1957:
1956:
1950:
1949:
1944:
1938:
1937:
1932:
1931:
1926:
1922:
1920:
1916:
1910:
1900:
1898:
1894:
1890:
1887:
1883:
1882:holocephalian
1879:
1875:
1871:
1866:
1865:
1860:
1856:
1852:
1849:
1845:
1841:
1840:Stensioellida
1835:
1834:
1829:
1823:Stensioellida
1820:
1818:
1815:
1811:
1807:
1804:
1800:
1796:
1792:
1788:
1784:
1783:
1778:
1774:
1773:
1768:
1764:
1758:
1757:
1752:
1743:
1741:
1740:
1734:
1730:
1729:
1724:
1720:
1714:
1713:
1708:
1699:
1697:
1693:
1689:
1688:chimaera-like
1685:
1679:
1678:
1673:
1664:
1662:
1658:
1653:
1649:
1645:
1641:
1635:
1634:
1629:
1620:
1618:
1614:
1610:
1606:
1601:
1597:
1593:
1589:
1585:
1584:Ptyctodontida
1579:
1575:
1569:Ptyctodontida
1566:
1564:
1560:
1556:
1550:
1549:
1544:
1535:
1533:
1532:
1527:
1523:
1522:
1517:
1513:
1510:
1506:
1502:
1501:Brindabella's
1498:
1497:
1490:
1489:
1484:
1475:
1473:
1472:
1467:
1466:
1461:
1457:
1453:
1452:pectoral fins
1449:
1444:
1440:
1436:
1432:
1428:
1422:
1421:
1416:
1407:
1405:
1404:
1399:
1398:
1393:
1392:
1387:
1386:
1381:
1377:
1376:Compagopiscis
1373:
1372:Compagopiscis
1369:
1365:
1361:
1357:
1351:
1350:
1345:
1339:
1338:
1333:
1319:
1317:
1314:
1310:
1306:
1305:
1300:
1296:
1286:
1284:
1283:osteostracans
1279:
1277:
1272:
1270:
1266:
1262:
1258:
1254:
1250:
1246:
1241:
1239:
1235:
1231:
1230:Louis Agassiz
1221:
1219:
1218:Carboniferous
1215:
1211:
1207:
1203:
1198:
1196:
1190:
1188:
1184:
1180:
1179:
1178:Xiushanosteus
1174:
1173:
1169:
1165:
1161:
1157:
1153:
1149:
1148:
1139:
1138:Entelognathus
1135:
1134:Entelognathus
1131:
1130:
1126:
1123:
1119:
1115:
1114:
1109:
1108:
1107:Entelognathus
1104:
1101:
1097:
1093:
1092:
1087:
1086:
1085:
1082:
1081:
1076:
1075:
1070:
1069:
1068:Entelognathus
1064:
1063:
1054:
1051:stage of the
1050:
1046:
1045:
1044:Xiushanosteus
1040:
1036:
1034:
1030:
1026:
1022:
1015:Fossil record
1012:
1010:
1006:
1005:Late Devonian
1002:
998:
992:
984:
980:
976:
975:
974:Eczematolepis
971:Fin spine of
969:
962:
961:Late Devonian
958:
954:
951:
947:
946:
941:
934:
930:
927:
923:
922:
917:
910:
906:
901:
892:
890:
886:
885:
879:
876:
872:
868:
864:
859:
857:
856:
851:
850:
845:
841:
840:
835:
831:
827:
826:
807:
803:
798:
796:
795:
790:
786:
785:
780:
776:
775:
770:
766:
762:
759:
755:
751:
750:
745:
741:
740:
735:
734:
729:
725:
723:
714:
713:
706:
702:
700:
696:
695:pulp cavities
692:
691:
690:Compagopiscis
686:
682:
678:
674:
670:
669:
664:
660:
656:
652:
648:
638:
636:
635:Carboniferous
632:
628:
624:
621:open for the
620:
616:
612:
611:
606:
605:
600:
599:
598:Eastmanosteus
594:
590:
586:
583:
579:
575:
571:
567:
563:
562:fossil record
558:
556:
552:
548:
544:
540:
535:
533:
529:
528:
523:
522:
517:
516:
511:
507:
504:precursor to
503:
499:
495:
491:
487:
483:
479:
475:
471:
466:
464:
460:
456:
452:
448:
444:
443:cartilaginous
440:
439:endoskeletons
436:
433:
429:
425:
422:
418:
414:
410:
407:
403:
400:
396:
393:' and δέρμα (
392:
388:
384:
380:
369:
368:
366:
363:
359:
354:
353:
348:
346:
342:
340:
339:
334:
332:
331:
330:Xiushanosteus
326:
322:
318:
316:
315:
310:
308:
307:
302:
300:
299:
294:
292:
291:
290:Entelognathus
286:
284:
283:
278:
277:
275:
272:
268:
264:
262:
258:
256:
255:Ptyctodontida
252:
250:
246:
243:
239:
235:
233:
230:
227:
226:
224:
219:
214:
209:
203:
200:
199:
196:
195:Gnathostomata
193:
191:Infraphylum:
190:
189:
186:
183:
180:
179:
176:
173:
170:
169:
166:
163:
160:
159:
156:
153:
150:
149:
144:
139:
135:
132:
131:pectoral fins
129:
125:
124:
118:
114:
109:
105:
101:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
39:
33:
22:
19:
6019:
5935:spiny sharks
5894:Halecomorphi
5713:Xenacanthida
5598:Phyllolepida
5564:
5531:Pituriaspida
5492:Ostracoderms
5426:Cyclostomata
5416:Jawless fish
5365:
5346:
5335:
5326:
5304:
5282:
5265:
5161:
5154:
5147:
5140:
5133:
5126:
5110:
5103:
5096:
5089:
5018:
5011:
5004:
4997:
4990:
4983:
4959:
4952:
4945:
4940:Ptyctodopsis
4938:
4931:
4924:
4917:
4912:Kimbryanodus
4910:
4903:
4896:
4889:
4882:
4875:
4870:Chelyophorus
4868:
4863:Campbellodus
4861:
4854:
4826:
4819:
4812:
4805:
4798:
4791:
4784:
4777:
4757:
4750:
4745:Eurycaraspis
4743:
4723:
4716:
4679:
4672:
4652:
4645:
4638:
4631:
4624:
4617:
4610:
4603:
4596:
4589:
4566:
4530:
4529:
4504:
4424:
4386:
4368:the original
4311:
4307:
4262:
4258:
4217:
4213:
4188:
4184:
4169:
4161:
4157:
4144:
4140:
4121:
4120:Janvier, P.
4081:
4077:
4026:
4022:
4016:
3965:
3959:
3942:
3909:
3905:
3901:
3895:
3881:
3872:
3862:
3844:
3825:
3812:
3803:
3799:
3793:
3781:. Retrieved
3770:
3721:
3717:
3707:
3696:the original
3681:
3677:
3664:
3632:(1): 41–51.
3629:
3625:
3615:
3606:
3597:
3586:. Retrieved
3582:the original
3572:
3560:
3548:
3536:
3524:
3497:
3493:
3483:
3456:
3452:
3442:
3430:
3385:
3381:
3371:
3354:
3350:
3314:
3310:
3300:
3257:
3253:
3243:
3231:. Retrieved
3177:
3173:
3154:
3147:
3121:paraphyletic
3115:
2950:Osteichthyes
2948:
2928:
2895:
2894:
2871:
2870:
2847:
2846:
2815:
2800:
2771:
2749:
2724:
2702:paraphyletic
2699:
2578:Titanichthys
2576:
2575:
2559:
2558:
2542:
2541:
2527:Brachyosteus
2525:
2524:
2503:Dunkleosteus
2501:
2500:
2479:Plourdosteus
2477:
2476:
2442:
2441:
2425:
2424:
2392:
2391:
2377:Pholidosteus
2375:
2374:
2326:
2312:Antineosteus
2310:
2309:
2293:
2292:
2260:
2240:Phyllolepida
2238:
2216:
2200:
2199:
2165:
2150:
2112:
2098:Yunnanolepis
2096:
2095:
2066:
2035:
2019:
2018:
1995:
1994:
1982:
1968:
1967:
1946:
1934:
1928:
1918:
1912:
1877:
1862:
1854:
1838:
1831:
1799:Phyllolepida
1780:
1770:
1761:
1754:
1737:
1732:
1726:
1717:
1710:
1682:
1675:
1638:
1631:
1582:
1577:
1555:Phyllolepida
1553:
1546:
1538:Phyllolepida
1529:
1526:jawless fish
1519:
1494:
1493:
1486:
1471:Bothriolepis
1469:
1465:Yunnanolepis
1463:
1425:
1418:
1403:Incisoscutum
1401:
1395:
1391:Dunkleosteus
1389:
1383:
1380:ichthyosaurs
1375:
1371:
1354:
1347:
1337:Dunkleosteus
1335:
1316:lagerstatten
1302:
1292:
1280:
1273:
1245:Erik Stensiö
1242:
1234:ostracoderms
1227:
1199:
1191:
1176:
1170:
1168:yunnanolepid
1164:Shimenolepis
1163:
1147:Shimenolepis
1145:
1143:
1137:
1133:
1127:
1111:
1105:
1096:Ludlow epoch
1089:
1078:
1072:
1066:
1060:
1058:
1042:
1018:
994:
972:
943:
921:Dunkleosteus
919:
882:
880:
860:
853:
847:
837:
823:
799:
792:
784:Titanichthys
782:
779:planktivores
772:
747:
737:
733:Fallacosteus
731:
722:Dunkleosteus
720:
718:
710:
688:
668:Bothriolepis
666:
655:Phyllolepida
644:
623:osteichthyan
610:Dunkleosteus
608:
602:
596:
559:
555:monophyletic
539:paraphyletic
536:
525:
519:
515:Incisoscutum
513:
467:
408:
394:
386:
382:
378:
377:
370:Placodermata
350:
336:
328:
312:
304:
296:
288:
280:
271:paraphyletic
242:Phyllolepida
207:
128:caliper-like
126:showing its
121:
18:
6044:Wikispecies
5955:lobe-finned
5881:Neopterygii
5867:Chondrostei
5803:Rhipidistia
5782:Lobe-finned
5732:Holocephali
5537:Osteostraci
5439:Hyperoartia
5284:Cathaymyrus
5252:Forerunners
5156:Stensioella
5112:Silurolepis
4992:Bolivosteus
4985:Asterosteus
4933:Palaeomylus
4919:Materpiscis
4884:Denisonodus
4598:Dobrowlania
4568:Hagiangella
4454:Subphylum:
4191:: 255–279.
4168:Long, J.A.
3091:Ostracoderm
3073:Fish portal
2394:Tapinosteus
2114:Euantiarcha
1970:Stensioella
1936:Pterichthys
1878:Stensioella
1855:Stensioella
1795:Arthrodires
1787:ptyctodonts
1661:arthrodires
1633:Asterosteus
1304:Stensioella
1276:gnathostome
1172:Zhanjilepis
1094:, from the
1091:Silurolepis
1062:Silurolepis
1029:arthrodires
977:, from the
945:Amazichthys
881:A study on
794:Heterosteus
789:Homostiidae
765:arthrodires
749:Amazichthys
681:arthrodires
633:during the
578:eurypterids
551:vertebrates
547:sister taxa
521:Materpiscis
498:paired fins
494:pelvic fins
492:to develop
482:gill arches
441:are mainly
426:during the
419:known from
352:Stensioella
298:Silurolepis
181:Subphylum:
6126:Placoderms
6120:Categories
6050:Placodermi
6020:Placodermi
5940:placoderms
5899:Ginglymodi
5833:Ray-finned
5798:Coelacanth
5643:Acanthodii
5580:Arthrodira
5565:Placoderms
5553:Jawed fish
5525:Galeaspida
5507:Thelodonti
5306:Haikouella
5297:Olfactores
5065:Arthrodira
5054:Arthrodira
4999:Gemuendina
4961:Rhynchodus
4926:Meeksiella
4905:Goniosteus
4877:Ctenurella
4828:Wijdeaspis
4619:Kosoraspis
4612:Kolymaspis
4591:Arabosteus
4531:Placodermi
4514:see below↓
4505:Placodermi
4456:Vertebrata
4425:Placodermi
4375:Placoderms
4164:: 263–308.
3588:2014-06-27
3459:(3): 318.
3135:References
3081:Acanthodii
2934:Acanthodii
2897:Janusiscus
2802:Arthrodira
2427:Coccosteus
2188:Arthrodira
1983:Placodermi
1907:See also:
1864:Gemuendina
1842:(" little
1806:trichotomy
1775:, and the
1733:Wijdeaspis
1559:stargazers
1458:-like, or
1431:Arthrodira
1397:Rolfosteus
1385:Arctolepis
1366:-nibbling
1356:Arthrodira
1349:Coccosteus
1327:Arthrodira
1202:freshwater
1160:Ludfordian
1156:Llandovery
1080:Wangolepis
999:and early
989:See also:
953:arthrodire
926:vertebrate
905:placoderms
884:Kolymaspis
863:homologous
855:Cowralepis
834:viviparous
754:morphology
739:Rolfosteus
728:vertebrate
604:Dinichthys
589:orthocones
582:cephalopod
574:arthropods
532:live birth
502:homologous
470:jawed fish
409:Placodermi
399:vertebrate
379:Placoderms
240:(includes
238:Arthrodira
208:Placodermi
185:Vertebrata
120:Fossil of
100:Llandovery
5998:† extinct
5853:Cladistia
5610:Rhenanida
5574:Antiarchi
5479:Conodonta
5458:Conodonts
5042:Antiarchi
5031:Antiarchi
5020:Ohioaspis
4976:Rhenanida
4947:Ptyctodus
4654:Tlamaspis
4640:Romundina
4442:Kingdom:
4295:205215898
4250:205213348
4114:205217467
3990:1476-4687
3762:252569910
3746:1476-4687
3656:1866-3508
3565:Long 1984
3516:2296-701X
3475:1424-2818
3453:Diversity
3435:Long 1983
3292:235477130
3140:Citations
2751:Antiarchi
2444:Torosteus
2068:Rhenanida
1930:Homostius
1915:cladogram
1903:Cladogram
1870:rhenanids
1817:dichotomy
1814:Rhenanida
1803:Antiarchi
1777:Rhenanida
1692:chimaeras
1657:antiarchs
1652:predators
1640:Rhenanida
1623:Rhenanida
1596:shellfish
1588:chimaeras
1460:arthropod
1427:Antiarchi
1410:Antiarchi
1299:monotypic
1253:Stockholm
1247:, at the
1214:Famennian
1206:saltwater
1189:, China.
1187:Chongqing
1183:Telychian
1122:tetrapods
1118:bony fish
1049:Telychian
1025:antiarchs
997:bony fish
983:Wisconsin
955:from the
875:hox genes
806:Kimberley
761:organisms
677:substrate
673:predators
659:Antiarchi
647:Rhenanida
543:outgroups
510:tetrapods
506:hindlimbs
421:Paleozoic
415:group of
261:Rhenanida
232:Antiarchi
161:Kingdom:
155:Eukaryota
24:Placoderm
6029:Wikidata
5816:Lungfish
5761:Chimaera
5513:Anaspida
5312:Tunicate
5290:Lancelet
5185:Category
5142:Minjinia
5013:Nefudina
5006:Jagorina
4793:Lunaspis
4647:Sudaspis
4633:Radotina
4605:Kimaspis
4450:Chordata
4448:Phylum:
4444:Animalia
4338:22219394
4287:19242474
4242:18509443
4106:19597477
4059:45922669
4051:27846567
4008:25581798
3934:45258255
3926:19215000
3873:Academia
3754:36171378
3692:19733953
3422:20479258
3333:27920231
3284:34146483
3202:23075852
3045:See also
2295:Holonema
1921:(2009):
1893:bodyplan
1889:Devonian
1851:Hunsrück
1808:and the
1767:Hunsrück
1728:Lunaspis
1712:Lunaspis
1696:rostrums
1648:ray-like
1617:claspers
1605:shagreen
1563:flatfish
1443:eye-hole
1439:tunicate
1364:detritus
1313:Hunsruck
1210:Frasnian
1195:Devonian
1021:Silurian
933:predator
774:Holonema
724:terrelli
685:nektonic
631:radiated
595:such as
585:molluscs
570:dominant
500:and the
445:, their
432:Devonian
430:and the
428:Silurian
362:Synonyms
314:Minjinia
175:Chordata
171:Phylum:
165:Animalia
151:Domain:
104:Devonian
6090:5292584
6064:4655826
6035:Q130932
5971:Related
5904:Teleost
5444:Lamprey
5434:Hagfish
5105:Qilinyu
4329:3367742
4267:Bibcode
4222:Bibcode
4193:Bibcode
4086:Bibcode
4031:Bibcode
4023:Science
3999:5536226
3970:Bibcode
3783:18 July
3726:Bibcode
3634:Bibcode
3413:2890420
3390:Bibcode
3262:Bibcode
3233:May 30,
3210:4302415
3182:Bibcode
2849:Qilinyu
1859:ratfish
1844:Stensio
1609:dentine
1456:caliper
1448:sinuous
1129:Qilinyu
1074:Qilinyu
959:of the
950:pelagic
911:, 2005.
844:clasper
817:⁄
758:pelagic
752:, with
699:dentine
572:marine
478:evolved
476:likely
472:(their
463:species
435:periods
424:fossils
413:extinct
404:of the
402:animals
282:Qilinyu
221:Orders
201:Class:
102:– Late
5267:Pikaia
5076:Others
4336:
4326:
4293:
4285:
4259:Nature
4248:
4240:
4214:Nature
4176:
4128:
4112:
4104:
4078:Nature
4057:
4049:
4006:
3996:
3988:
3961:Nature
3932:
3924:
3832:
3806:: 304.
3779:. UCMP
3760:
3752:
3744:
3718:Nature
3689:
3654:
3514:
3473:
3420:
3410:
3331:
3290:
3282:
3208:
3200:
3174:Nature
1919:et al.
1295:orders
1265:sharks
1257:sharks
1238:turtle
1100:Yunnan
1001:sharks
791:, and
657:, and
619:niches
587:(e.g.
580:) and
576:(e.g.
459:scaled
451:thorax
387:plakos
215:, 1848
6103:34249
6072:IRMNG
5923:Lists
5723:Shark
4291:S2CID
4246:S2CID
4110:S2CID
4055:S2CID
3930:S2CID
3822:(PDF)
3758:S2CID
3699:(PDF)
3686:INIST
3674:(PDF)
3288:S2CID
3206:S2CID
3107:Notes
1644:Rhine
1152:Hunan
1033:basal
663:genus
490:clade
486:teeth
411:, an
406:class
395:derma
391:plate
213:McCoy
98:Late
6077:1495
5950:bony
5773:Bony
5237:Fish
5063:see
5040:see
4491:see
4334:PMID
4283:PMID
4238:PMID
4174:ISBN
4126:ISBN
4102:PMID
4047:PMID
4004:PMID
3986:ISSN
3922:PMID
3830:ISBN
3785:2011
3750:PMID
3742:ISSN
3652:ISSN
3512:ISSN
3471:ISSN
3418:PMID
3329:PMID
3280:PMID
3235:2008
3198:PMID
1933:and
1731:and
1659:and
1204:and
1120:and
1071:and
1027:and
1007:and
948:, a
852:and
736:and
625:and
524:and
474:jaws
449:and
447:head
383:plax
38:PreꞒ
6059:EoL
5718:Ray
5322:? (
4324:PMC
4316:doi
4275:doi
4263:457
4230:doi
4218:453
4201:doi
4162:107
4094:doi
4082:460
4039:doi
4027:354
3994:PMC
3978:doi
3966:520
3914:doi
3910:270
3804:248
3734:doi
3722:609
3642:doi
3502:doi
3461:doi
3408:PMC
3398:doi
3386:107
3359:doi
3319:doi
3270:doi
3190:doi
3178:491
3119:If
1561:or
1362:to
1261:wax
1251:in
1098:of
981:of
545:or
508:in
389:) '
6122::
6100::
6087::
6074::
6061::
6046::
6031::
5656:†"
5331:,
4332:.
4322:.
4310:.
4306:.
4289:.
4281:.
4273:.
4261:.
4244:.
4236:.
4228:.
4216:.
4199:.
4189:38
4187:.
4160:.
4156:.
4145:26
4143:.
4139:.
4108:.
4100:.
4092:.
4080:.
4053:.
4045:.
4037:.
4025:.
4002:.
3992:.
3984:.
3976:.
3964:.
3958:.
3928:.
3920:.
3908:.
3871:.
3852:.
3802:.
3756:.
3748:.
3740:.
3732:.
3720:.
3716:.
3682:21
3680:.
3676:.
3650:.
3640:.
3630:15
3628:.
3624:.
3510:.
3500:.
3498:10
3496:.
3492:.
3469:.
3457:15
3455:.
3451:.
3416:.
3406:.
3396:.
3384:.
3380:.
3355:19
3353:.
3341:^
3327:.
3315:66
3313:.
3309:.
3286:.
3278:.
3268:.
3258:31
3256:.
3252:.
3218:^
3204:.
3196:.
3188:.
3176:.
3164:^
1819:.
1812:+
1801:+
1797:+
1642:("
1499:("
1318:.
1271:.
1220:.
1197:.
1162:.
1055:.
1011:.
804:,
797:.
665:,
653:,
649:,
637:.
601:,
557:.
534:.
518:,
465:.
385:,
349:?†
343:?†
88:Pg
32:Ma
5886:†
5847:"
5843:"
5841:†
5808:†
5790:†
5754:†
5748:†
5742:†
5736:†
5711:†
5705:†
5699:†
5693:†
5669:†
5663:†
5660:"
5650:†
5640:†
5631:?
5627:†
5624:?
5620:†
5614:†
5608:†
5602:†
5596:†
5590:†
5584:†
5578:†
5572:†
5562:†
5535:†
5529:†
5523:†
5517:†
5511:†
5505:†
5499:†
5489:†
5477:†
5471:†
5468:?
5464:†
5456:†
5351:?
5344:†
5340:)
5333:†
5324:†
5317:†
5302:†
5280:†
5263:†
5218:e
5211:t
5204:v
5115:?
4582:*
4555:*
4427:"
4423:"
4416:e
4409:t
4402:v
4340:.
4318::
4312:8
4297:.
4277::
4269::
4252:.
4232::
4224::
4207:.
4203::
4195::
4116:.
4096::
4088::
4061:.
4041::
4033::
4010:.
3980::
3972::
3936:.
3916::
3889:.
3875:.
3838:.
3787:.
3764:.
3736::
3728::
3658:.
3644::
3636::
3591:.
3567:.
3555:.
3543:.
3531:.
3518:.
3504::
3477:.
3463::
3437:.
3424:.
3400::
3392::
3365:.
3361::
3335:.
3321::
3294:.
3272::
3264::
3237:.
3212:.
3192::
3184::
1212:–
1124:.
985:.
963:.
935:.
819:2
815:1
812:+
810:2
335:†
327:†
319:?
311:†
303:†
295:†
287:†
279:†
273:)
269:(
265:†
259:†
253:†
247:†
244:)
236:†
229:†
205:†
93:N
83:K
78:J
73:T
68:P
63:C
58:D
53:S
48:O
43:Ꞓ
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
