3495:
3500:
104:
125:
418:, and that the mouth-plates retained the prey while allowing the water to escape as it closed its mouth. A study has since confirmed this assumption as its jaws are functionally closer to that of filter feeders like baleen whales and basking sharks, and it appears to have developed from benthic durophagists that became pelagic suspension feeders. This would make it the first (known) large-sized vertebrate filter feeder.
1018:
2742:
2730:
2716:
2723:
3490:
2736:
984:
This is the type species, from the
Cleveland Shale. Its infrognathals are strongly recurved medially, and is elongated with a spatula-like process at the anterior end. The headshield averages about 60 cm (24 in) in length.
1013:
s. The cranial roof is comparatively narrower and more rounded. It is the largest known species in the genus, and possibly one of the largest
Devonian vertebrates known. The head is about 90 cm (35 in) in length.
1035:
This
Cleveland Shale species is known from a badly preserved, incomplete infragnathal. It was originally described by Claypole as "Brontichthys clarki" in 1894. As "Brontichthys" is a junior synonym of
1078:
This species placement within the genus is in doubt. It is based on incomplete nuchal and central plates found in Upper
Famennian-aged marine strata of the Holy Cross Mountains in Poland.
1088:
This species is found in Lower
Famennian-aged marine strata of Tafilalet, Southern Morocco. The fossil material of this species strongly suggests it is as large as the Cleveland Shale'
1287:
3700:
3592:
3858:
3782:
3687:
3579:
3868:
3494:
1444:
1231:
1148:
3863:
3833:
1547:
995:
This
Cleveland Shale species is based on an infragnathal bone more than 36 cm (14 in) in length. May possibly be a synonym of
1519:
1485:
1264:
3843:
3838:
3499:
1302:
1502:(2018) "New information on Titanichthys (Placodermi, Arthrodira) from the Cleveland Shale (Upper Devonian) of Ohio, USA"
124:
3848:
3828:
571:
3773:
1364:"New information on Titanichthys (Placodermi, Arthrodira) from the Cleveland Shale (Upper Devonian) of Ohio, USA"
1110:- that is, after Termier was able to convince his colleagues that the bone scraps were of a placoderm, and not a
1540:
554:
523:
475:
2741:
771:
3853:
1990:
604:
422:
was estimated to have reached a length of 7โ7.6 m (23โ25 ft), but
Engelman (2023) suggested that
3519:
2729:
2722:
1630:
3795:
3726:
3618:
3489:
822:
710:
2307:
2881:
3823:
3674:
3665:
3566:
3272:
1375:
1177:
971:
686:
3279:
2438:
2368:
756:
453:
3230:
2964:
2715:
2533:
2323:
2079:
1657:
1533:
838:
795:
499:
2805:
2654:
2184:
2004:
398:
had small, ineffective-looking mouth-plates that lacked a sharp cutting edge. It is assumed that
2798:
2526:
2519:
2254:
2238:
2114:
1401:
1281:
1068:' infragnathal is much straighter, and does not have a spatula-like process on its anterior end.
628:
343:
119:
3800:
3731:
3118:
3066:
2940:
2668:
2452:
2424:
2403:
3623:
3557:
3265:
2925:
2833:
2735:
2473:
2431:
2417:
2282:
1825:
3713:
3605:
3597:
2316:
2011:
1931:
1811:
1797:
1515:
1481:
1440:
1393:
1327:"A Devonian Fish Tale: A New Method of Body Length Estimation Suggests Much Smaller Sizes for
1260:
1227:
1203:
1144:
3718:
3705:
3610:
3053:
2647:
1871:
1383:
1342:
1193:
1185:
652:
103:
3258:
3189:
3139:
3084:
2912:
2847:
2771:
2759:
2679:
2612:
2382:
2156:
2027:
1922:
967:
808:
665:
442:
208:
1379:
1181:
3396:
3304:
3202:
3132:
3111:
2990:
2633:
2589:
2560:
2544:
2504:
2410:
2268:
2195:
2056:
1938:
1880:
1804:
1198:
1165:
1047:
1042:
3817:
3431:
3403:
3389:
3375:
3347:
3338:
3237:
3209:
3151:
3125:
3104:
2948:
2933:
2865:
2780:
2640:
2394:
2339:
2226:
2177:
2142:
2045:
1981:
1860:
1615:
1592:
1501:
1405:
1106:
1101:
970:, though species are also found in similarly aged strata in Morocco and possibly the
726:
403:
195:
61:
3466:
3361:
3313:
3097:
3073:
3013:
3006:
2999:
2979:
2956:
2896:
2854:
2840:
2789:
2661:
2605:
2553:
2497:
2466:
2353:
2261:
2247:
2211:
2170:
2121:
2100:
2093:
2086:
1997:
1898:
1832:
1818:
1776:
1762:
1755:
390:
251:
3650:
3542:
1254:
1221:
3767:
3659:
3551:
3452:
3424:
3410:
3368:
3354:
3327:
3286:
3167:
3020:
2889:
2619:
2598:
2484:
2296:
2275:
2163:
1952:
1769:
1164:
Coatham, Samuel J.; Vinther, Jakob; Rayfield, Emily J.; Klug, Christian (2020).
415:
36:
1304:
Fishes, Living and Fossil: An
Outline of Their Forms and Probable Relationships
3459:
3445:
3438:
3417:
3382:
3244:
3223:
3216:
3027:
2874:
2826:
2819:
2812:
2574:
2218:
2204:
2149:
2128:
2107:
2065:
1959:
1945:
1911:
1887:
1853:
1839:
1790:
1723:
1710:
1669:
1586:
1556:
447:
388:, Eastern North America, and possibly Europe. Many of the species approached
182:
81:
46:
3758:
1397:
3320:
3251:
3174:
3160:
3034:
2972:
2626:
2567:
2511:
2459:
2360:
2289:
2135:
2072:
1783:
1748:
1741:
1642:
1598:
963:
377:
169:
136:
86:
1207:
1009:
This
Cleveland Shale species has infragnathals that are not as recurved as
430:, likely measuring about or just over 4.1 metres (13.5 ft) in length.
1017:
3752:
3644:
3536:
2445:
2375:
1966:
1846:
1580:
1388:
1363:
1111:
394:
in size and build. Unlike its relative, however, the various species of
381:
156:
76:
71:
56:
51:
41:
3787:
1189:
3692:
3584:
1096:
is estimated to be 200 cm (79 in) When the first fossils of
407:
385:
370:
91:
66:
1347:
1326:
1040:, it should not be confused with another, similarly-named arthrodire,
1574:
1060:
This
Cleveland Shale species has an infragnathal as large as that of
406:
that used its capacious mouth to swallow or inhale schools of small,
146:
3513:
1016:
411:
373:
3571:
1092:. The average combined length of the head and trunk shields for
3679:
3517:
2757:
1708:
1567:
1529:
1166:"Was the Devonian placoderm Titanichthys a suspension feeder?"
1525:
1253:
Charlie Underwood, Martha Richter, Zerina Johanson (2019).
1439:. Stuttgart New York: Gustav Fischer Verlag. p. 101.
1226:. Ohio. Division of Geological Survey. 1996. p. 290.
1143:. Stuttgart New York: Gustav Fischer Verlag. p. 100.
1104:, the specimens were originally placed within the genus
3742:
3634:
3526:
3337:
3303:
3188:
3150:
3083:
3052:
2989:
2911:
2864:
2788:
2779:
2770:
2678:
2588:
2543:
2483:
2393:
2337:
2306:
2237:
2194:
2055:
2025:
1980:
1921:
1897:
1870:
1721:
1668:
1641:
1614:
1511:The Rise of Fishes: 500 Million Years of Evolution
1479:. Oxford University Press, Oxford, New York.
1541:
1513:. Johns Hopkins University Press, Baltimore.
1362:Boyle, James; Ryan, Michael J. (March 2017).
8:
1286:: CS1 maint: multiple names: authors list (
3514:
3194:
3089:
3058:
2917:
2785:
2776:
2767:
2754:
2489:
2345:
2037:
2033:
1903:
1733:
1729:
1718:
1705:
1647:
1620:
1611:
1564:
1548:
1534:
1526:
1427:
1425:
1423:
1421:
1419:
1417:
1415:
1259:. Cambridge University Press. p. 13.
1131:
1129:
1127:
102:
20:
1436:Volume 2 of Handbook of Paleoichthyology'
1387:
1346:
1197:
1140:Volume 2 of Handbook of Paleoichthyology'
962:The genus shows a great diversity in the
1123:
1279:
1459:See Janvier (1998) p.323 for details.
7:
1307:. Macmillan and Company. p. 130
1256:Evolution and Development of Fishes
814:
787:
763:
702:
678:
671:
644:
620:
596:
546:
539:
515:
491:
467:
460:
14:
3859:Famennian genus first appearances
3498:
3493:
3488:
2740:
2734:
2728:
2721:
2714:
441:is thought to have been a basal
123:
1499:James Boyle and Michael J. Ryan
380:from shallow seas of the Late
1:
3869:Fossil taxa described in 1885
1325:Engelman, Russell K. (2023).
3864:Famennian genus extinctions
3834:Placoderms of North America
111:Rendered reconstruction of
3885:
1170:Royal Society Open Science
572:Kendrickichthys cavernosus
426:was comparable in size to
376:of giant, aberrant marine
16:Extinct genus of placoderm
3486:
3197:
3092:
3061:
2920:
2766:
2753:
2712:
2492:
2348:
2040:
2036:
1906:
1736:
1732:
1717:
1704:
1650:
1623:
1610:
1563:
1331:(Placodermi: Arthrodira)"
836:
819:
812:
792:
785:
768:
761:
724:
707:
700:
683:
676:
669:
649:
642:
625:
618:
601:
594:
568:
551:
544:
537:
520:
513:
496:
489:
472:
465:
349:
342:
276:
271:
257:
250:
120:Scientific classification
118:
110:
101:
23:
1432:Denison, Robert (1978).
1136:Denison, Robert (1978).
1100:were found by geologist
555:Bruntonichthys multidens
524:Trematosteus fontanellus
476:Rhachiosteus pterygiatus
410:-like fish, or possibly
305:Wright in Claypole, 1893
1368:Journal of Paleontology
772:Tafilalichthys lavocati
605:Bullerichthys fascidens
1301:Bashford Dean (1895).
1025:
1021:Fossil head shield of
3796:Paleobiology Database
3774:Titanichthys agassizi
3744:Titanichthys agassizi
3727:Paleobiology Database
3619:Paleobiology Database
1329:Dunkleosteus terrelli
1020:
823:Titanichthys agassizi
711:Heintzichthys gouldii
445:, closest related to
261:Titanichthys agassizi
3844:Placoderms of Africa
3839:Paleontology in Ohio
1389:10.1017/jpa.2016.136
1023:Titanichthys clarkii
972:Holy Cross Mountains
687:Gorgonichthys clarki
113:Titanichthys clarkii
1380:2017JPal...91..318B
1190:10.1098/rsos.200272
1182:2020RSOS....700272C
796:Bungartius perissus
500:Tapinosteus heintzi
3849:Fossils of Morocco
2239:Groenlandaspididae
1026:
629:Dinichthys herzeri
3829:Arthrodire genera
3811:
3810:
3714:Open Tree of Life
3606:Open Tree of Life
3520:Taxon identifiers
3511:
3510:
3507:
3506:
3484:
3483:
3480:
3479:
3476:
3475:
3184:
3183:
3048:
3047:
3044:
3043:
2907:
2906:
2749:
2748:
2710:
2709:
2706:
2705:
2702:
2701:
2698:
2697:
2584:
2583:
2333:
2332:
2308:Williamsaspididae
2021:
2020:
1976:
1975:
1932:Austrophyllolepis
1798:Heightingtonaspis
1700:
1699:
1696:
1695:
1692:
1691:
1688:
1687:
1522:
1488:
1477:Early Vertebrates
1473:Janvier, Philippe
1446:978-0-89574-027-4
1348:10.3390/d15030318
1233:978-0-931079-05-4
1150:978-0-89574-027-4
954:
953:
945:
944:
936:
935:
927:
926:
918:
917:
909:
908:
900:
899:
891:
890:
882:
881:
873:
872:
864:
863:
855:
854:
746:
745:
737:
736:
583:
582:
362:
361:
357:
336:
326:
316:
306:
296:
286:
246:
227:
3876:
3804:
3803:
3791:
3790:
3778:
3777:
3776:
3763:
3762:
3761:
3735:
3734:
3722:
3721:
3709:
3708:
3696:
3695:
3683:
3682:
3670:
3669:
3668:
3655:
3654:
3653:
3627:
3626:
3614:
3613:
3601:
3600:
3588:
3587:
3575:
3574:
3562:
3561:
3560:
3547:
3546:
3545:
3515:
3502:
3497:
3492:
3195:
3090:
3059:
3054:Pachyosteomorphi
2918:
2882:Kimberleyichthys
2786:
2777:
2768:
2755:
2744:
2738:
2733:
2732:
2726:
2725:
2718:
2648:Parabuchanosteus
2490:
2346:
2038:
2034:
1904:
1872:Wuttagoonaspidae
1734:
1730:
1719:
1706:
1648:
1621:
1612:
1605:
1604:
1565:
1550:
1543:
1536:
1527:
1514:
1508:
1500:
1480:
1474:
1460:
1457:
1451:
1450:
1429:
1410:
1409:
1391:
1359:
1353:
1352:
1350:
1322:
1316:
1315:
1313:
1312:
1298:
1292:
1291:
1285:
1277:
1275:
1273:
1250:
1244:
1243:
1241:
1240:
1218:
1212:
1211:
1201:
1161:
1155:
1154:
1133:
815:
788:
764:
703:
679:
672:
653:Hadrosteus rapax
645:
621:
597:
547:
540:
516:
492:
468:
461:
355:
334:
324:
314:
304:
294:
284:
244:
237:
225:
220:
207:
194:
181:
168:
128:
127:
106:
96:
33:
29:Temporal range:
21:
3884:
3883:
3879:
3878:
3877:
3875:
3874:
3873:
3814:
3813:
3812:
3807:
3799:
3794:
3786:
3781:
3772:
3771:
3766:
3757:
3756:
3751:
3738:
3730:
3725:
3717:
3712:
3704:
3699:
3691:
3686:
3678:
3673:
3666:Titanichthyidae
3664:
3663:
3658:
3649:
3648:
3643:
3636:Titanichthyidae
3630:
3622:
3617:
3609:
3604:
3596:
3591:
3583:
3578:
3570:
3565:
3556:
3555:
3550:
3541:
3540:
3535:
3522:
3512:
3503:
3472:
3333:
3299:
3273:Squamatognathus
3259:Kendrickichthys
3190:Aspinothoracidi
3180:
3146:
3140:Xiangshuiosteus
3085:Dunkleosteoidea
3079:
3040:
2985:
2913:Incisoscutoidea
2903:
2860:
2848:Protitanichthys
2772:Coccosteomorphi
2762:
2760:Eubrachythoraci
2745:
2727:
2720:
2694:
2680:Eubrachythoraci
2674:
2613:Burrinjucosteus
2580:
2539:
2479:
2389:
2383:Xiangshuiosteus
2342:
2329:
2302:
2233:
2190:
2157:Neophlyctaenius
2051:
2030:
2028:Phlyctaenioidei
2017:
1972:
1923:Phyllolepididae
1917:
1893:
1866:
1726:
1713:
1684:
1664:
1637:
1606:
1570:
1569:
1559:
1554:
1506:
1498:
1495:
1493:Further reading
1472:
1469:
1464:
1463:
1458:
1454:
1447:
1431:
1430:
1413:
1361:
1360:
1356:
1324:
1323:
1319:
1310:
1308:
1300:
1299:
1295:
1278:
1271:
1269:
1267:
1252:
1251:
1247:
1238:
1236:
1234:
1220:
1219:
1215:
1163:
1162:
1158:
1151:
1135:
1134:
1125:
1120:
1086:
1076:
1058:
1033:
1007:
993:
982:
968:Cleveland Shale
960:
955:
946:
937:
928:
919:
910:
901:
892:
883:
874:
865:
856:
809:Titanichthyidae
747:
738:
666:Aspinothoracida
584:
436:
354:
335:Kulczycki, 1957
267:
264:
243:
235:
224:
222:Titanichthyidae
218:
209:Dinichthyloidea
205:
192:
179:
166:
122:
97:
95:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
39:
31:
30:
27:
17:
12:
11:
5:
3882:
3880:
3872:
3871:
3866:
3861:
3856:
3854:Famennian life
3851:
3846:
3841:
3836:
3831:
3826:
3816:
3815:
3809:
3808:
3806:
3805:
3792:
3779:
3764:
3748:
3746:
3740:
3739:
3737:
3736:
3723:
3710:
3697:
3684:
3671:
3656:
3640:
3638:
3632:
3631:
3629:
3628:
3615:
3602:
3589:
3576:
3563:
3548:
3532:
3530:
3524:
3523:
3518:
3509:
3508:
3505:
3504:
3487:
3485:
3482:
3481:
3478:
3477:
3474:
3473:
3471:
3470:
3463:
3456:
3449:
3442:
3435:
3428:
3421:
3414:
3407:
3400:
3397:Gymnotrachelus
3393:
3386:
3379:
3372:
3365:
3358:
3351:
3343:
3341:
3335:
3334:
3332:
3331:
3324:
3317:
3309:
3307:
3305:Brachydeiridae
3301:
3300:
3298:
3297:
3290:
3283:
3280:Tafilalichthys
3276:
3269:
3262:
3255:
3248:
3241:
3234:
3227:
3220:
3213:
3206:
3203:Bruntonichthys
3198:
3192:
3186:
3185:
3182:
3181:
3179:
3178:
3171:
3164:
3156:
3154:
3148:
3147:
3145:
3144:
3136:
3133:Westralichthys
3129:
3122:
3115:
3112:Golshanichthys
3108:
3101:
3093:
3087:
3081:
3080:
3078:
3077:
3070:
3062:
3056:
3050:
3049:
3046:
3045:
3042:
3041:
3039:
3038:
3031:
3024:
3017:
3010:
3003:
2995:
2993:
2991:Camuropiscidae
2987:
2986:
2984:
2983:
2976:
2969:
2961:
2953:
2945:
2937:
2930:
2921:
2915:
2909:
2908:
2905:
2904:
2902:
2901:
2893:
2886:
2878:
2870:
2868:
2862:
2861:
2859:
2858:
2851:
2844:
2837:
2830:
2823:
2816:
2809:
2802:
2794:
2792:
2783:
2774:
2764:
2763:
2758:
2751:
2750:
2747:
2746:
2713:
2711:
2708:
2707:
2704:
2703:
2700:
2699:
2696:
2695:
2693:
2692:
2691:
2690:
2684:
2682:
2676:
2675:
2673:
2672:
2665:
2658:
2651:
2644:
2637:
2634:Goodradigbeeon
2630:
2623:
2616:
2609:
2602:
2594:
2592:
2590:Buchanosteidae
2586:
2585:
2582:
2581:
2579:
2578:
2571:
2564:
2561:Atlantidosteus
2557:
2549:
2547:
2545:Migmatocephala
2541:
2540:
2538:
2537:
2530:
2523:
2516:
2508:
2505:Cathlesichthys
2501:
2493:
2487:
2481:
2480:
2478:
2477:
2470:
2463:
2456:
2449:
2442:
2439:Gyroplacosteus
2435:
2428:
2421:
2414:
2411:Belemnacanthus
2407:
2399:
2397:
2391:
2390:
2388:
2387:
2379:
2372:
2369:Gemuendenaspis
2365:
2357:
2349:
2343:
2338:
2335:
2334:
2331:
2330:
2328:
2327:
2320:
2312:
2310:
2304:
2303:
2301:
2300:
2293:
2286:
2279:
2272:
2269:Groenlandaspis
2265:
2258:
2251:
2243:
2241:
2235:
2234:
2232:
2231:
2223:
2215:
2208:
2200:
2198:
2196:Arctolepididae
2192:
2191:
2189:
2188:
2181:
2174:
2167:
2160:
2153:
2146:
2139:
2132:
2125:
2118:
2111:
2104:
2097:
2090:
2083:
2076:
2069:
2061:
2059:
2057:Phlyctaeniidae
2053:
2052:
2050:
2049:
2041:
2031:
2026:
2023:
2022:
2019:
2018:
2016:
2015:
2008:
2001:
1994:
1986:
1984:
1978:
1977:
1974:
1973:
1971:
1970:
1963:
1956:
1949:
1942:
1939:Cobandrahlepis
1935:
1927:
1925:
1919:
1918:
1916:
1915:
1907:
1901:
1895:
1894:
1892:
1891:
1884:
1881:Wuttagoonaspis
1876:
1874:
1868:
1867:
1865:
1864:
1857:
1850:
1843:
1836:
1829:
1822:
1815:
1808:
1805:Kujdanowiaspis
1801:
1794:
1787:
1780:
1773:
1766:
1759:
1752:
1745:
1737:
1727:
1722:
1715:
1714:
1709:
1702:
1701:
1698:
1697:
1694:
1693:
1690:
1689:
1686:
1685:
1683:
1682:
1681:
1680:
1674:
1672:
1666:
1665:
1663:
1662:
1661:
1660:
1651:
1645:
1639:
1638:
1636:
1635:
1634:
1633:
1624:
1618:
1608:
1607:
1603:
1602:
1595:
1589:
1583:
1577:
1568:
1561:
1560:
1555:
1553:
1552:
1545:
1538:
1530:
1524:
1523:
1504:
1494:
1491:
1490:
1489:
1468:
1465:
1462:
1461:
1452:
1445:
1411:
1374:(2): 318โ336.
1354:
1317:
1293:
1265:
1245:
1232:
1213:
1176:(5): 200โ272.
1156:
1149:
1122:
1121:
1119:
1116:
1085:
1080:
1075:
1070:
1057:
1052:
1048:Dunkleosteidae
1043:Bruntonichthys
1032:
1027:
1006:
1001:
992:
987:
981:
976:
959:
956:
952:
951:
948:
947:
943:
942:
939:
938:
934:
933:
930:
929:
925:
924:
921:
920:
916:
915:
912:
911:
907:
906:
903:
902:
898:
897:
894:
893:
889:
888:
885:
884:
880:
879:
876:
875:
871:
870:
867:
866:
862:
861:
858:
857:
853:
852:
849:
848:
835:
832:
831:
828:
827:
818:
813:
811:
805:
804:
801:
800:
791:
786:
784:
781:
780:
777:
776:
767:
762:
760:
757:Mylostomatidae
753:
752:
749:
748:
744:
743:
740:
739:
735:
734:
731:
730:
723:
720:
719:
716:
715:
706:
701:
699:
696:
695:
692:
691:
682:
677:
675:
670:
668:
662:
661:
658:
657:
648:
643:
641:
638:
637:
634:
633:
624:
619:
617:
614:
613:
610:
609:
600:
595:
593:
590:
589:
586:
585:
581:
580:
577:
576:
567:
564:
563:
560:
559:
550:
545:
543:
538:
536:
533:
532:
529:
528:
519:
514:
512:
509:
508:
505:
504:
495:
490:
488:
485:
484:
481:
480:
471:
466:
464:
459:
454:Tafilalichthys
443:aspinothoracid
435:
432:
360:
359:
356:Claypole, 1894
347:
346:
340:
339:
338:
337:
331:T. ?koslowskii
327:
317:
307:
297:
295:Newberry, 1887
287:
285:Newberry, 1887
274:
273:
269:
268:
266:Newberry, 1885
265:
255:
254:
248:
247:
245:Newberry, 1885
233:
229:
228:
216:
212:
211:
203:
199:
198:
190:
186:
185:
177:
173:
172:
164:
160:
159:
154:
150:
149:
144:
140:
139:
134:
130:
129:
116:
115:
108:
107:
99:
98:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
40:
35:
34:
28:
15:
13:
10:
9:
6:
4:
3:
2:
3881:
3870:
3867:
3865:
3862:
3860:
3857:
3855:
3852:
3850:
3847:
3845:
3842:
3840:
3837:
3835:
3832:
3830:
3827:
3825:
3822:
3821:
3819:
3802:
3797:
3793:
3789:
3784:
3780:
3775:
3769:
3765:
3760:
3754:
3750:
3749:
3747:
3745:
3741:
3733:
3728:
3724:
3720:
3715:
3711:
3707:
3702:
3698:
3694:
3689:
3685:
3681:
3676:
3672:
3667:
3661:
3657:
3652:
3646:
3642:
3641:
3639:
3637:
3633:
3625:
3620:
3616:
3612:
3607:
3603:
3599:
3594:
3590:
3586:
3581:
3577:
3573:
3568:
3564:
3559:
3553:
3549:
3544:
3538:
3534:
3533:
3531:
3529:
3525:
3521:
3516:
3501:
3496:
3491:
3469:
3468:
3464:
3462:
3461:
3457:
3455:
3454:
3450:
3448:
3447:
3443:
3441:
3440:
3436:
3434:
3433:
3432:Paramylostoma
3429:
3427:
3426:
3422:
3420:
3419:
3415:
3413:
3412:
3408:
3406:
3405:
3404:Heintzichthys
3401:
3399:
3398:
3394:
3392:
3391:
3390:Gorgonichthys
3387:
3385:
3384:
3380:
3378:
3377:
3376:Draconichthys
3373:
3371:
3370:
3366:
3364:
3363:
3359:
3357:
3356:
3352:
3350:
3349:
3348:Alienacanthus
3345:
3344:
3342:
3340:
3339:Selenosteidae
3336:
3330:
3329:
3325:
3323:
3322:
3318:
3316:
3315:
3311:
3310:
3308:
3306:
3302:
3296:
3295:
3291:
3289:
3288:
3284:
3282:
3281:
3277:
3275:
3274:
3270:
3268:
3267:
3263:
3261:
3260:
3256:
3254:
3253:
3249:
3247:
3246:
3242:
3240:
3239:
3238:Erromenosteus
3235:
3233:
3232:
3231:Dinomylostoma
3228:
3226:
3225:
3221:
3219:
3218:
3214:
3212:
3211:
3210:Bullerichthys
3207:
3205:
3204:
3200:
3199:
3196:
3193:
3191:
3187:
3177:
3176:
3172:
3170:
3169:
3165:
3163:
3162:
3158:
3157:
3155:
3153:
3152:Heterosteidae
3149:
3142:
3141:
3137:
3135:
3134:
3130:
3128:
3127:
3126:Kiangyousteus
3123:
3121:
3120:
3116:
3114:
3113:
3109:
3107:
3106:
3105:Eastmanosteus
3102:
3100:
3099:
3095:
3094:
3091:
3088:
3086:
3082:
3076:
3075:
3071:
3069:
3068:
3064:
3063:
3060:
3057:
3055:
3051:
3037:
3036:
3032:
3030:
3029:
3025:
3023:
3022:
3018:
3016:
3015:
3011:
3009:
3008:
3004:
3002:
3001:
2997:
2996:
2994:
2992:
2988:
2982:
2981:
2977:
2975:
2974:
2970:
2967:
2966:
2965:Parabelosteus
2962:
2959:
2958:
2954:
2951:
2950:
2949:Harrytoombsia
2946:
2943:
2942:
2938:
2936:
2935:
2934:Compagopiscis
2931:
2928:
2927:
2923:
2922:
2919:
2916:
2914:
2910:
2899:
2898:
2894:
2892:
2891:
2887:
2884:
2883:
2879:
2877:
2876:
2872:
2871:
2869:
2867:
2866:Panxiosteidae
2863:
2857:
2856:
2852:
2850:
2849:
2845:
2843:
2842:
2838:
2836:
2835:
2831:
2829:
2828:
2824:
2822:
2821:
2817:
2815:
2814:
2810:
2808:
2807:
2803:
2801:
2800:
2796:
2795:
2793:
2791:
2787:
2784:
2782:
2781:Coccosteoidea
2778:
2775:
2773:
2769:
2765:
2761:
2756:
2752:
2743:
2739:
2737:
2731:
2724:
2717:
2688:
2687:
2686:
2685:
2683:
2681:
2677:
2671:
2670:
2666:
2664:
2663:
2659:
2657:
2656:
2652:
2650:
2649:
2645:
2643:
2642:
2641:Narrominaspis
2638:
2636:
2635:
2631:
2629:
2628:
2624:
2622:
2621:
2617:
2615:
2614:
2610:
2608:
2607:
2603:
2601:
2600:
2596:
2595:
2593:
2591:
2587:
2577:
2576:
2572:
2570:
2569:
2565:
2563:
2562:
2558:
2556:
2555:
2551:
2550:
2548:
2546:
2542:
2536:
2535:
2534:Luetkeichthys
2531:
2529:
2528:
2524:
2522:
2521:
2517:
2514:
2513:
2509:
2507:
2506:
2502:
2500:
2499:
2495:
2494:
2491:
2488:
2486:
2482:
2476:
2475:
2471:
2469:
2468:
2464:
2462:
2461:
2457:
2455:
2454:
2450:
2448:
2447:
2443:
2441:
2440:
2436:
2434:
2433:
2429:
2427:
2426:
2422:
2420:
2419:
2415:
2413:
2412:
2408:
2406:
2405:
2401:
2400:
2398:
2396:
2395:Holonematidae
2392:
2385:
2384:
2380:
2378:
2377:
2373:
2371:
2370:
2366:
2363:
2362:
2358:
2356:
2355:
2351:
2350:
2347:
2344:
2341:
2340:Brachythoraci
2336:
2326:
2325:
2324:Williamsaspis
2321:
2319:
2318:
2314:
2313:
2311:
2309:
2305:
2299:
2298:
2294:
2292:
2291:
2287:
2285:
2284:
2280:
2278:
2277:
2273:
2271:
2270:
2266:
2264:
2263:
2259:
2257:
2256:
2252:
2250:
2249:
2245:
2244:
2242:
2240:
2236:
2229:
2228:
2227:Kueichowlepis
2224:
2221:
2220:
2216:
2214:
2213:
2209:
2207:
2206:
2202:
2201:
2199:
2197:
2193:
2187:
2186:
2182:
2180:
2179:
2178:Prosphymaspis
2175:
2173:
2172:
2168:
2166:
2165:
2161:
2159:
2158:
2154:
2152:
2151:
2147:
2145:
2144:
2143:Kueichowlepis
2140:
2138:
2137:
2133:
2131:
2130:
2126:
2124:
2123:
2119:
2117:
2116:
2112:
2110:
2109:
2105:
2103:
2102:
2098:
2096:
2095:
2091:
2089:
2088:
2084:
2082:
2081:
2080:Barwickosteus
2077:
2075:
2074:
2070:
2068:
2067:
2063:
2062:
2060:
2058:
2054:
2048:
2047:
2046:Dicksonosteus
2043:
2042:
2039:
2035:
2032:
2029:
2024:
2014:
2013:
2009:
2007:
2006:
2002:
2000:
1999:
1995:
1993:
1992:
1988:
1987:
1985:
1983:
1982:Actinolepidae
1979:
1969:
1968:
1964:
1962:
1961:
1957:
1955:
1954:
1950:
1948:
1947:
1943:
1941:
1940:
1936:
1934:
1933:
1929:
1928:
1926:
1924:
1920:
1914:
1913:
1909:
1908:
1905:
1902:
1900:
1896:
1890:
1889:
1885:
1883:
1882:
1878:
1877:
1875:
1873:
1869:
1863:
1862:
1861:Yujiangolepis
1858:
1856:
1855:
1851:
1849:
1848:
1844:
1842:
1841:
1837:
1835:
1834:
1830:
1828:
1827:
1823:
1821:
1820:
1816:
1814:
1813:
1809:
1807:
1806:
1802:
1800:
1799:
1795:
1793:
1792:
1788:
1786:
1785:
1781:
1779:
1778:
1774:
1772:
1771:
1767:
1765:
1764:
1760:
1758:
1757:
1753:
1751:
1750:
1746:
1744:
1743:
1739:
1738:
1735:
1731:
1728:
1725:
1720:
1716:
1712:
1707:
1703:
1678:
1677:
1676:
1675:
1673:
1671:
1667:
1659:
1655:
1654:
1653:
1652:
1649:
1646:
1644:
1640:
1632:
1631:Gnathostomata
1628:
1627:
1626:
1625:
1622:
1619:
1617:
1616:Gnathostomata
1613:
1609:
1600:
1596:
1594:
1593:Gnathostomata
1591:Infraphylum:
1590:
1588:
1584:
1582:
1578:
1576:
1572:
1571:
1566:
1562:
1558:
1551:
1546:
1544:
1539:
1537:
1532:
1531:
1528:
1521:
1520:0-8018-5438-5
1517:
1512:
1507:Long, John A.
1505:
1503:
1497:
1496:
1492:
1487:
1486:0-19-854047-7
1483:
1478:
1471:
1470:
1466:
1456:
1453:
1448:
1442:
1438:
1435:
1428:
1426:
1424:
1422:
1420:
1418:
1416:
1412:
1407:
1403:
1399:
1395:
1390:
1385:
1381:
1377:
1373:
1369:
1365:
1358:
1355:
1349:
1344:
1340:
1336:
1332:
1330:
1321:
1318:
1306:
1305:
1297:
1294:
1289:
1283:
1268:
1266:9781107179448
1262:
1258:
1257:
1249:
1246:
1235:
1229:
1225:
1224:
1217:
1214:
1209:
1205:
1200:
1195:
1191:
1187:
1183:
1179:
1175:
1171:
1167:
1160:
1157:
1152:
1146:
1142:
1139:
1132:
1130:
1128:
1124:
1117:
1115:
1113:
1109:
1108:
1107:Gorgonichthys
1103:
1102:Henri Termier
1099:
1095:
1091:
1084:
1081:
1079:
1074:
1073:T. kozlowskii
1071:
1069:
1067:
1063:
1056:
1053:
1051:
1049:
1045:
1044:
1039:
1031:
1028:
1024:
1019:
1015:
1012:
1005:
1002:
1000:
998:
991:
990:T. attenuatus
988:
986:
980:
977:
975:
973:
969:
965:
957:
950:
949:
941:
940:
932:
931:
923:
922:
914:
913:
905:
904:
896:
895:
887:
886:
878:
877:
869:
868:
860:
859:
851:
850:
847:
846:
845:
841:
834:
833:
830:
829:
826:
825:
824:
817:
816:
810:
807:
806:
803:
802:
799:
798:
797:
790:
789:
783:
782:
779:
778:
775:
774:
773:
766:
765:
758:
755:
754:
751:
750:
742:
741:
733:
732:
729:
728:
727:Selenosteidae
722:
721:
718:
717:
714:
713:
712:
705:
704:
698:
697:
694:
693:
690:
689:
688:
681:
680:
674:
673:
667:
664:
663:
660:
659:
656:
655:
654:
647:
646:
640:
639:
636:
635:
632:
631:
630:
623:
622:
616:
615:
612:
611:
608:
607:
606:
599:
598:
592:
591:
588:
587:
579:
578:
575:
574:
573:
566:
565:
562:
561:
558:
557:
556:
549:
548:
542:
541:
535:
534:
531:
530:
527:
526:
525:
518:
517:
511:
510:
507:
506:
503:
502:
501:
494:
493:
487:
486:
483:
482:
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478:
477:
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469:
463:
462:
458:
456:
455:
450:
449:
444:
440:
433:
431:
429:
425:
421:
417:
413:
409:
405:
404:filter feeder
401:
397:
393:
392:
387:
383:
379:
375:
372:
368:
367:
358:
353:
348:
345:
341:
333:
332:
328:
323:
322:
318:
313:
312:
308:
303:
302:
301:T. attenuatus
298:
293:
292:
288:
283:
282:
278:
277:
275:
270:
263:
262:
256:
253:
249:
242:
241:
234:
231:
230:
223:
217:
214:
213:
210:
204:
202:Superfamily:
201:
200:
197:
196:Brachythoraci
191:
188:
187:
184:
178:
175:
174:
171:
165:
162:
161:
158:
155:
152:
151:
148:
145:
142:
141:
138:
135:
132:
131:
126:
121:
117:
114:
109:
105:
100:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
38:
26:
22:
19:
3743:
3635:
3558:Titanichthys
3528:Titanichthys
3527:
3467:Walterosteus
3465:
3458:
3451:
3444:
3437:
3430:
3423:
3416:
3409:
3402:
3395:
3388:
3381:
3374:
3367:
3362:Brachyosteus
3360:
3353:
3346:
3326:
3319:
3314:Brachydeirus
3312:
3294:Titanichthys
3293:
3292:
3285:
3278:
3271:
3264:
3257:
3250:
3243:
3236:
3229:
3222:
3215:
3208:
3201:
3173:
3166:
3159:
3138:
3131:
3124:
3117:
3110:
3103:
3098:Dunkleosteus
3096:
3074:Rhachiosteus
3072:
3065:
3033:
3026:
3019:
3014:Incisoscutum
3012:
3007:Fallacosteus
3005:
3000:Camuropiscis
2998:
2980:Trematosteus
2978:
2971:
2963:
2957:Mcnamaraspis
2955:
2947:
2939:
2932:
2924:
2897:Plourdosteus
2895:
2888:
2880:
2873:
2855:Watsonosteus
2853:
2846:
2841:Millerosteus
2839:
2832:
2825:
2818:
2811:
2806:Clarkeosteus
2804:
2797:
2790:Coccosteidae
2719:
2667:
2662:Taemasosteus
2660:
2655:Toombsosteus
2653:
2646:
2639:
2632:
2625:
2618:
2611:
2606:Buchanosteus
2604:
2597:
2573:
2566:
2559:
2554:Antineosteus
2552:
2532:
2525:
2518:
2510:
2503:
2498:Angarichthys
2496:
2472:
2467:Tropidosteus
2465:
2458:
2451:
2444:
2437:
2430:
2423:
2416:
2409:
2402:
2381:
2374:
2367:
2359:
2354:Antineosteus
2352:
2322:
2315:
2295:
2288:
2281:
2274:
2267:
2262:Cosmacanthus
2260:
2253:
2248:Africanaspis
2246:
2225:
2217:
2212:Heintzosteus
2210:
2203:
2185:Stipatosteus
2183:
2176:
2171:Phlyctaenius
2169:
2162:
2155:
2148:
2141:
2134:
2127:
2122:Heterogaspis
2120:
2113:
2106:
2101:Elegantaspis
2099:
2094:Diadsomaspis
2092:
2087:Cartieraspis
2085:
2078:
2071:
2064:
2044:
2010:
2005:Bollandaspis
2003:
1998:Ailuracantha
1996:
1989:
1965:
1958:
1951:
1944:
1937:
1930:
1910:
1899:Phyllolepida
1886:
1879:
1859:
1852:
1845:
1838:
1833:Proaethaspis
1831:
1824:
1819:Lehmanosteus
1817:
1810:
1803:
1796:
1789:
1782:
1777:Bryantolepis
1775:
1768:
1763:Antarctaspis
1761:
1756:Anarthraspis
1754:
1747:
1740:
1510:
1476:
1467:Bibliography
1455:
1437:
1433:
1371:
1367:
1357:
1338:
1334:
1328:
1320:
1309:. Retrieved
1303:
1296:
1270:. Retrieved
1255:
1248:
1237:. Retrieved
1222:
1216:
1173:
1169:
1159:
1141:
1137:
1105:
1097:
1093:
1089:
1087:
1082:
1077:
1072:
1065:
1061:
1059:
1054:
1041:
1038:Titanichthys
1037:
1034:
1030:T. hussakofi
1029:
1022:
1011:T. agassizi'
1010:
1008:
1003:
996:
994:
989:
983:
978:
961:
843:
840:Titanichthys
839:
837:
821:
820:
794:
793:
770:
769:
725:
709:
708:
685:
684:
651:
650:
627:
626:
603:
602:
570:
569:
553:
552:
522:
521:
498:
497:
474:
473:
452:
446:
439:Titanichthys
438:
437:
428:Dunkleosteus
427:
424:Titanichthys
423:
420:Titanichthys
419:
400:Titanichthys
399:
396:Titanichthys
395:
391:Dunkleosteus
389:
366:Titanichthys
365:
364:
363:
352:Brontichthys
351:
350:
330:
329:
325:Lehman, 1954
320:
319:
311:T. hussakofi
310:
309:
300:
299:
290:
289:
280:
279:
260:
259:
252:Type species
240:Titanichthys
239:
238:
221:
112:
25:Titanichthys
24:
18:
3824:Arthrodires
3768:Wikispecies
3660:Wikispecies
3552:Wikispecies
3453:Selenosteus
3425:Pachyosteus
3411:Melanosteus
3369:Braunosteus
3355:Amazichthys
3328:Synauchenia
3287:Tapinosteus
3168:Heterosteus
3021:Latocamurus
2890:Panxiosteus
2799:Belgiosteus
2620:Errolosteus
2599:Arenipiscis
2527:Euleptaspis
2520:Euleptaspis
2485:Homostiidae
2297:Turrisaspis
2276:Mithakaspis
2255:Boomeraspis
2164:Pageauaspis
2115:Grifftaylor
1991:Actinolepis
1953:Phyllolepis
1770:Baringaspis
1585:Subphylum:
1434:Placodermi
1272:1 September
1223:Bulletin 70
1138:Placodermi
1098:T. termieri
1094:T. termieri
1083:T. termieri
997:T. agassizi
979:T. agassizi
974:in Poland.
416:zooplankton
321:T. termieri
281:T. agassizi
3818:Categories
3460:Stenosteus
3446:Rhinosteus
3439:Phymosteus
3418:Microsteus
3383:Enseosteus
3245:Hadrosteus
3224:Dinichthys
3217:Bungartius
3119:Hussakofia
3067:Leptosteus
3028:Rolfosteus
2941:Cyrtosteus
2875:Janiosteus
2827:Jiuchengia
2820:Dickosteus
2813:Coccosteus
2689:see belowโ
2669:Uralosteus
2575:Tityosteus
2453:Megaloplax
2425:Deirosteus
2404:Artesonema
2219:Huginaspis
2205:Arctolepis
2150:Lyhoalepis
2129:Huginaspis
2108:Gaspeaspis
2066:Aggeraspis
1960:Placolepis
1946:Cowralepis
1912:Gavinaspis
1888:Yiminaspis
1854:Simblaspis
1840:Qataraspis
1791:Eskimaspis
1724:Arthrodira
1711:Arthrodira
1679:see belowโ
1670:Arthrodira
1658:Placodermi
1643:Placodermi
1599:Placodermi
1587:Vertebrata
1557:Arthrodira
1341:(3). 318.
1311:2022-08-28
1239:2022-08-28
1118:References
1090:T. clarkii
1062:T. clarkii
1004:T. clarkii
448:Bungartius
226:Dean, 1901
189:Suborder:
183:Arthrodira
170:Placodermi
3651:Q18602551
3543:Q18602550
3321:Oxyosteus
3266:Mylostoma
3252:Holdenius
3175:Yinostius
3161:Herasmius
3035:Tubonasus
2973:Torosteus
2926:Belosteus
2834:Livosteus
2627:Exutaspis
2568:Homosteus
2512:Dhanguura
2474:Tropinema
2460:Rhenonema
2432:Deveonema
2418:Bimbianga
2361:Dhanguura
2290:Tiaraspis
2283:Mulgaspis
2136:Kolpaspis
2073:Arctaspis
1826:Mediaspis
1784:Erikaspis
1749:Aleosteus
1742:Aethaspis
1573:Kingdom:
1406:132831650
1398:0022-3360
1335:Diversity
1282:cite book
1066:T. rectus
1064:, though
1055:T. rectus
964:Famennian
434:Phylogeny
378:placoderm
315:Hay, 1930
291:T. clarki
143:Kingdom:
137:Eukaryota
32:Famennian
3788:11174219
3753:Wikidata
3645:Wikidata
3572:57245902
3537:Wikidata
2446:Holonema
2376:Maideria
2317:Elvaspis
2012:Szelepis
1967:Yurammia
1847:Sigaspis
1812:Lataspis
1597:Class: "
1581:Chordata
1579:Phylum:
1575:Animalia
1509:(1996):
1475:(1998):
1208:32537223
1112:dinosaur
382:Devonian
344:Synonyms
272:Species
215:Family:
157:Chordata
153:Phylum:
147:Animalia
133:Domain:
3759:Q517892
3719:4139753
3693:4853026
3680:4655856
3611:5292599
3598:1035664
3585:7237315
1376:Bibcode
1199:7277245
1178:Bibcode
958:Species
408:anchovy
386:Morocco
371:extinct
232:Genus:
176:Order:
163:Class:
3801:390786
3732:352344
3706:101233
1518:
1484:
1443:
1404:
1396:
1263:
1230:
1206:
1196:
1147:
966:-aged
844:clarki
414:-like
402:was a
369:is an
3701:IRMNG
3624:34397
3593:IRMNG
1402:S2CID
412:krill
374:genus
3783:GBIF
3688:GBIF
3580:GBIF
1656:see
1629:see
1516:ISBN
1482:ISBN
1441:ISBN
1394:ISSN
1288:link
1274:2022
1261:ISBN
1228:ISBN
1204:PMID
1145:ISBN
842:cf.
451:and
37:Pre๊
3675:EoL
3567:EoL
1384:doi
1343:doi
1194:PMC
1186:doi
1046:of
384:of
3820::
3798::
3785::
3770::
3755::
3729::
3716::
3703::
3690::
3677::
3662::
3647::
3621::
3608::
3595::
3582::
3569::
3554::
3539::
1414:^
1400:.
1392:.
1382:.
1372:91
1370:.
1366:.
1339:15
1337:.
1333:.
1284:}}
1280:{{
1202:.
1192:.
1184:.
1172:.
1168:.
1126:^
1114:.
1050:.
999:.
759:?
457:.
87:Pg
3143:?
2968:?
2960:?
2952:?
2944:?
2929:?
2900:?
2885:?
2515:?
2386:?
2364:?
2230:?
2222:?
1601:"
1549:e
1542:t
1535:v
1449:.
1408:.
1386::
1378::
1351:.
1345::
1314:.
1290:)
1276:.
1242:.
1210:.
1188::
1180::
1174:7
1153:.
258:โ
236:โ
219:โ
206:โ
193:โ
180:โ
167:โ
92:N
82:K
77:J
72:T
67:P
62:C
57:D
52:S
47:O
42:๊
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