1126:
1204:
7662:
6429:
7797:
5231:
1327:
143:
7809:
5237:
919:
4614:
6445:
7669:
4619:
4073:
7675:
4081:
508:
4088:
118:
6437:
1153:, the two front teeth pointed forward and were higher on the jaw than the other teeth, while the next three pairs of teeth pointed to the sides. The final two (preserved) pairs of teeth pointed downward. Finally, a unique oval depression was located below the first pair of teeth.
2846:
Borja
Holgado, Rodrigo V. Pêgas, José Ignacio Canudo, Josep Fortuny, Taissa Rodrigues, Julio Company & Alexander W.A. Kellner, 2019, "On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria",
1189:, which may be its closest relative or a member of the same genus. It also had a straight, rather than curved, front margin, unlike its relatives, and begins at the tip of the snout, rather than further back as in other species.
2332:
Kellner, Alexander W. A.; Campos, Diogenes A.; Sayão, Juliana M.; Saraiva, Antônio A.F.; Rodrigues, Taissa; Oliveira, Gustavo; Cruz, Lilian A.; Costa, Fabiana R.; Silva, Helder P.; Ferreira, Jennyfer S. (March 2013).
1239:
from other species. The jaw tip is nearly 10 cm tall and 5.6 cm wide, with teeth up to 1.3 cm in base diameter. If the proportions of this specimen were consistent with other known species of
1183:
had a keel-shaped crest on the front of its jaws, though it was broad and thinned from base to top, rather than the uniformly thin crests of its relatives. This kind of thickened crest is also seen in
719:. Rodrigues and Kellner disagreed with this classification, however, noting that both did not possess the unique straightened crest beginning at the snout tip, or sideways pointed teeth, of
746:(such as a flattened upper surface of the snout), though Rodrigues and Kellner regarded them as dubious, and noted that they are also present in other genera. Kellner recombined it as
653:. The revival of the genus meant that of several related species, then assigned to other genera, had to be re-evaluated to determine whether or not they actually belonged to
1244:, the total skull length could have been up to 75 cm, leading to an estimated wingspan of 7 metres (23 ft). However, this species may belong to a different genus.
2786:
Pentland, Adele H.; Poropat, Stephen F.; Tischler, Travis R.; Sloan, Trish; Elliott, Robert A.; Elliott, Harry A.; Elliott, Judy A.; Elliott, David A. (December 2019).
1323:, more specifically within the subfamily Coloborhynchinae. The cladogram on the left shows a phylogenetic analysis made by Borja Holgado and Rodrigo Pêgas in 2020.
1168:, the tip of the snout flared out into a wider rosette, in contrast to the narrow posterior jaws. However, whereas the rosettes of species typically assigned to
2716:"First occurrence of the pterosaur Coloborhynchus (Pterosauria, Ornithocheiridae) from the Wessex Formation (Lower Cretaceous) of the Isle of Wight, England"
869:
530:
has a highly convoluted history of classification. Over the years numerous species have been assigned to it, and often, species have been shuffled between
2553:
Smith, Roy E.; Ibrahim, Nizar; Longrich, Nicholas; Unwin, David M.; Jacobs, Megan L.; Williams, Cariad J.; Zouhri, Samir; Martill, David M. (2023-02-04).
3095:
2934:
gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of
Cretaceous toothed pterodactyloids".
591:. Owen considered the defining trait of the genus to be the location of the front tooth pairs high on the side of the upper jaws. However, in 1913
2871:
Kellner, Alexander W. A.; Caldwell, Michael W.; Holgado, Borja; Vecchia, Fabio M. Dalla; Nohra, Roy; Sayão, Juliana M.; Currie, Philip J. (2019).
880:
based on a partial snout, and also tentatively referred another specimen from the same locality to a different, unnamed species. A 2020 review of
7754:
633:
7834:
705:
2343:
Wellnhofer, 1987 (Pterodactyloidea, Anhangueridae) and other large pterosaurs from the
Romualdo Formation, Lower Cretaceous, Brazil"
7741:
2634:
taxonomy revisited: is our understanding of
Santana Group pterosaur diversity biased by poor biological and stratigraphic control?"
1315:
in their analysis. They published their conclusion in 2019. However, many subsequent analyses made in 2019 and 2020 have recovered
2216:. However one study found it among the carnivorous pterosaur taxa, as opposed to several other piscivore and insectivore species.
2792:
gen. et sp. nov., a new ornithocheirid pterosaur from the Winton
Formation (Cenomanian–lower Turonian) of Queensland, Australia"
7844:
2715:
2683:
Martill, David M.; Unwin, David M. (April 2012). "The world's largest toothed pterosaur, NHMUK R481, an incomplete rostrum of
2470:
811:, the second of which being one of the original members of the genus according to Richard Owen in 1874. According to Kellner,
3074:
2969:
Myers, Timothy S. (2017). "Diet of ornithocheiroid pterosaurs inferred from stable carbon isotope analysis of tooth enamel".
910:
to be subtle enough to justify their synonymy, stating that the material was damaged and fragmentary enough to support this.
3048:
2522:"A taxonomic and phylogenetic review of the anhanguerid pterosaur group Coloborhynchinae and the new clade Tropeognathinae"
7839:
7661:
2307:
1235:
described in 2011 consists of a very large upper jaw tip which displays the tooth characteristics that distinguish
2994:"Dietary diversity and evolution of the earliest flying vertebrates revealed by dental microwear texture analysis"
1149:, it can only be differentiated from its relatives based on its unique combination of tooth socket positions. In
1125:
2973:. GSA Annual Meeting in Seattle, Washington, USA. Vol. 49. Geological Society of America. p. 305496.
2230:
601:, the second genus and species Owen erected in 1874. Hooley also ignored Owen's reassignment of the two former
5236:
1307:
were assigned to the
Ornithocheiridae, however, unlike the study made by Andres and Myers, they used the name
1145:
is known only from a partial upper jaw. Therefore, according to
Rodrigues and Kellner's 2008 re-evaluation on
661:, again based mainly on the unique positions of the tooth sockets. Rodrigues and Kellner argued that Lee's
142:
6428:
5230:
595:
concluded that this location was an artefact of the erosion and that the genus was indistinguishable from
571:. The genus name means "maimed beak", a reference to the damaged and eroded condition of the fossil; the
7787:
2602:
1018:
752:
489:
was once thought to be the largest known toothed pterosaur, however, a specimen of the closely related
2428:
792:, lacked its unique characteristics of the tooth row, a position also supported by Fastnacht in 2001.
762:
from the genus, based on lack of comparable diagnostic features. Unwin, in 2001, assigned the species
4613:
3067:
2398:
1192:
A second specimen showing all of these same unique features was reported to
Brazilian paleontologist
1087:
7829:
7732:
3171:
2225:
638:
4072:
4649:
4080:
2873:"First complete pterosaur from the Afro-Arabian continent: insight into pterodactyloid diversity"
572:
298:
4618:
2433:, with a review of the specimens from the Cambridge Greensand in the Sedgwick Museum, Cambridge"
7772:
7694:
7569:
5727:
2153:
1514:
1158:
1079:
1037:
970:
710:
680:
592:
7759:
7668:
477:
age, 140 to 136 million years ago), and depending on which species are included, possibly the
7813:
7767:
7674:
1203:
6436:
5822:
4261:
4171:
4087:
3060:
3005:
2992:
Bestwick, Jordan; Unwin, David M.; Butler, Richard J.; Purnell, Mark A. (28 October 2020).
2884:
2803:
6444:
8:
7801:
7326:
7015:
6702:
6001:
5928:
3696:
3416:
1228:
1073:
964:
796:
542:
279:
3009:
2888:
2807:
1227:
specimen), and indeed the largest toothed pterosaur known. A referred specimen from the
6855:
5329:
5172:
4315:
4118:
4105:
4019:
3861:
3492:
3467:
3028:
2993:
2951:
2907:
2872:
2824:
2787:
2765:
2660:
2629:
2497:
2283:
2256:
317:
137:
7281:
6096:
6025:
5267:
5254:
5197:
4960:
4858:
4760:
4678:
4456:
4433:
4399:
4329:
3870:
3786:
3738:
3712:
3425:
3330:
3033:
2955:
2912:
2829:
2665:
2576:
2501:
2364:
2288:
1193:
689:
647:
523:
2769:
2359:
2334:
827:(in fact it lacks a crest altogether), and may instead belong to the same genus as "
657:. In 2008, Taissa Rodrigues and Alexander Kellner re-formulated the key features of
7510:
7393:
7385:
7177:
7106:
7031:
6573:
6476:
6463:
6395:
6275:
6264:
5514:
5371:
5308:
5083:
5065:
4903:
4727:
4543:
4529:
4487:
3993:
3979:
3911:
3590:
3140:
3023:
3013:
2974:
2943:
2902:
2892:
2855:
2819:
2811:
2757:
2730:
2696:
2655:
2645:
2566:
2533:
2489:
2444:
2354:
2278:
2268:
1879:
1563:
1536:
1410:
1299:
also recovered a similar cladogram compared to Andres and Myers (2013), where both
1252:
1207:
881:
466:
458:
200:
30:
2947:
2700:
2493:
575:
means "key snout", referring to its form in cross-section. Owen also reclassified
7599:
7442:
7291:
7076:
6980:
6955:
6948:
6935:
6868:
6754:
6652:
6618:
6514:
6494:
6348:
6206:
6147:
6134:
6105:
5981:
5892:
5751:
5624:
5579:
5451:
5380:
5359:
5185:
4846:
4826:
4798:
4774:
4710:
4702:
4695:
4687:
4442:
4422:
4252:
4228:
4178:
3986:
3969:
3959:
3936:
3892:
3884:
3852:
3841:
3820:
3774:
3766:
3705:
3621:
3561:
3480:
3449:
3339:
3318:
3271:
3156:
3131:
2734:
2086:
1760:
1497:
902:. However, a 2023 review of Kem Kem pterosaurs found the traits that distinguish
758:
685:
2978:
7576:
7549:
7351:
7265:
7250:
7141:
7126:
7092:
6917:
6901:
6889:
6875:
6828:
6785:
6745:
6659:
6605:
6333:
6326:
6319:
6291:
6198:
6119:
6046:
5957:
5713:
5605:
5540:
5481:
5421:
5387:
5107:
5034:
5006:
4925:
4875:
4814:
4783:
4383:
4359:
4337:
4283:
4241:
4185:
4153:
3834:
3751:
3728:
3607:
3533:
3501:
3354:
3303:
3251:
3018:
2897:
2859:
2815:
2571:
2554:
2019:
1958:
1601:
1473:
1280:
1164:
1101:
1051:
1003:
984:
671:
669:, and from an earlier time period, belonged in its own genus, which they named
547:
447:
374:
216:
2761:
2750:
Earth and
Environmental Science Transactions of the Royal Society of Edinburgh
2448:
1326:
7823:
7717:
7613:
7584:
7560:
7541:
7499:
7484:
7433:
7422:
7315:
7307:
7257:
7194:
7165:
7149:
7133:
7119:
7060:
7038:
6989:
6882:
6643:
6586:
6505:
6340:
6312:
6284:
6155:
6038:
5829:
5759:
5631:
5614:
5470:
5407:
5351:
5116:
5072:
4953:
4946:
4939:
4932:
4917:
4867:
4791:
4748:
4741:
4667:
4514:
4507:
4500:
4349:
4291:
4276:
4269:
4217:
4000:
3669:
3629:
3579:
3548:
3509:
3404:
3396:
3374:
3278:
3262:
2580:
2273:
2127:
2110:
1998:
1798:
1781:
1750:
1550:
1483:
1460:
1320:
1223:
1185:
1138:
1065:
952:
873:
764:
560:
491:
454:
349:
68:
7606:
7534:
7405:
7300:
7234:
7186:
6996:
6924:
6802:
6770:
6538:
6305:
6233:
6174:
6069:
5706:
5691:
5675:
5666:
5655:
5495:
5458:
5344:
5294:
5137:
5050:
5043:
5023:
4995:
4988:
4974:
4967:
4882:
4839:
4806:
4522:
4464:
4407:
4392:
4372:
4197:
3928:
3920:
3900:
3827:
3806:
3799:
3654:
3442:
3388:
3347:
3037:
2916:
2833:
2748:
Andres, Brian; Myers, Timothy S. (September 2012). "Lone Star
Pterosaurs".
2669:
2538:
2521:
2368:
2292:
2053:
1941:
1427:
941:
894:
837:
605:
species, leaving them in that genus. In 1967, Kuhn agreed with Hooley that
597:
588:
538:
386:
287:
245:
236:
708:
supported this position, and also synonymized the more well-known species
7726:
7492:
7452:
7371:
7335:
7069:
6844:
6777:
6720:
6666:
6523:
6191:
6062:
6054:
6014:
5973:
5965:
5840:
5807:
5775:
5767:
5738:
5699:
5683:
5642:
5571:
5563:
5556:
5502:
5443:
5435:
5096:
5058:
4981:
4636:
4569:
4536:
4472:
3813:
3682:
3637:
3614:
3289:
2607:
sp. nov. (Pterodactyloidea) from the Albian (Lower Cretaceous) of Brazil"
1357:
564:
474:
43:
2475:(Pterosauria, Ornithocheiridae) from the mid-Cretaceous of North Africa"
2315:
1287:
recovered a similar result to the one by Andres and Myers in 2013 where
918:
7746:
7460:
7415:
7343:
7218:
7210:
7045:
7024:
6969:
6837:
6812:
6762:
6727:
6673:
6561:
6553:
6546:
6298:
6221:
6213:
6112:
6076:
5914:
5815:
5794:
5783:
5720:
5337:
5130:
5123:
4558:
4449:
3456:
3113:
2650:
1848:
1625:
898:, and also referred the unnamed Ifezouane Formation coloborhynchine to
511:
482:
88:
53:
2930:
Pêgas, Rodrigo V.; Holgado, Borja; Leal, Maria Eduarda C. (2021). "On
742:
by Veldmeijer in 2003. It shares one or two characters in common with
7520:
6962:
6910:
6632:
6625:
6593:
6531:
6257:
6250:
6242:
6162:
5940:
5906:
5856:
5849:
5586:
5524:
5428:
5016:
4415:
4132:
3759:
3599:
3434:
3311:
3233:
3220:
3183:
3122:
3083:
2213:
1893:
450:
187:
154:
93:
37:
7688:
1196:
by Darren Naish in 2007, and likely represents a second specimen of
7711:
7592:
7476:
7468:
7242:
7226:
7157:
6734:
6084:
5989:
5921:
5593:
5548:
5321:
4550:
4210:
3107:
2036:
1831:
1642:
1372:
621:
species. Most later researchers followed these opinions, regarding
556:
496:
174:
125:
83:
78:
63:
58:
48:
1279:, and the family itself is placed within the more inclusive clade
852:, and placed most other species in other genera, or declared them
819:, and its precise classification is open to debate. He noted that
1232:
877:
568:
507:
470:
117:
98:
73:
6183:
5950:
5532:
4140:
4007:
3101:
2469:
Jacobs, M.L.; Martill, D.M.; Ibrahim, N.; Longrich, N. (2019).
1263:
A topology made by Andres and Myers in 2013 placed the species
788:
as distinct, and noted that like the other species assigned to
693:
665:, which possessed some differences in the skull and teeth from
643:
478:
164:
3052:
2250:
2248:
2246:
1133:
holotype with dotted lines reconstructing worn away jaw margin
795:
In 2001, Unwin also reassigned the two other species from the
5281:
3877:
2870:
2335:"The largest flying reptile from Gondwana: a new specimen of
738:
Another Brazilian species from the Romualdo Member was named
650:
443:
2785:
2468:
1221:
represents the second largest known ornithocheirid (after a
2555:"The pterosaurs of the Cretaceous Kem Kem Group of Morocco"
2392:
2390:
2388:
2386:
2384:
2382:
2380:
2378:
2243:
2991:
2687:(Seeley, 1870) from the Cambridge Greensand of England".
2552:
2331:
617:
as the type species of the genus, rather than one of the
559:
BMNH 1822, a partial snout from the Hastings Beds of the
16:
Genus of anhanguerid pterosaur from the Cretaceous period
2375:
1295:. Their cladogram is shown on the left. In 2018, Jacobs
613:. Furthermore, Kuhn was the first to formally designate
587:, though he did not designate any of these three as the
1275:) within the family Ornithocheiridae as sister taxa to
922:
Holotype and referred specimen of the possible species
457:, though it has also been recovered as a member of the
4647:
3049:
The Pterosaur Database (multiple species, scroll down)
7785:
2676:
2628:
Pinheiro, Felipe L.; Rodrigues, Taissa (4 May 2017).
2261:
complex (Pterosauria) from the Cretaceous of England"
294:
275:
252:
2397:Rodrigues, Taiss; Kellner, Alexander W. A. (2008).
1106:Seeley, 1870 = "Ptenodactylus capito" Seeley, 1869
3695:
2741:
5171:
4116:
2627:
2327:
2325:
7821:
5265:
2929:
2396:
2254:
6474:
6394:
5307:
2781:
2779:
913:
723:. Instead, Rodrigues and Kellner regarded both
3155:
2923:
2322:
1114:Jacobs, Martill, Ibrahim & Longrich, 2018
933:by various scientists over the years include:
485:ages (113 to 93.9 million years ago) as well.
3068:
2515:
2513:
2511:
502:
7059:
2864:
2776:
2707:
2682:
2596:
2594:
2592:
2590:
2519:
2308:"'Ugly' fossil is largest toothed pterosaur"
6173:
2840:
2747:
784:as a distinct genus in 2008, also regarded
534:and related genera by various researchers.
3231:
3075:
3061:
2723:Proceedings of the Geologists' Association
2600:
2508:
2212:is traditionally thought to be an oceanic
1333:holotype with interpretative line drawings
844:In 2013, Rodrigues and Kellner considered
526:pterosaurs named during the 19th century,
116:
3027:
3017:
2906:
2896:
2823:
2659:
2649:
2587:
2570:
2537:
2358:
2282:
2272:
2399:"Review of the pterodactyloid pterosaur
2347:Anais da Academia Brasileira de Ciências
1325:
1202:
1200:, though it has not yet been described.
1172:were rounded and spoon-shaped, those of
1124:
917:
823:does not possess the unique features of
506:
2713:
2464:
2462:
2460:
2458:
7822:
4104:
2437:Annals and Magazine of Natural History
2426:
2305:
815:is too incomplete to fully compare to
7835:Early Cretaceous pterosaurs of Europe
7693:
7692:
6461:
5253:
5252:
4634:
4103:
3218:
3094:
3056:
2968:
2420:
750:in 2006. Similarly, Kellner excluded
678:A partial lower jaw originally named
6462:
2455:
1337:
929:Species which have been assigned to
495:is now thought to have had a larger
2429:"LXI.— On the Ornithosaurian genus
2255:Rodrigues, T.; Kellner, A. (2013).
13:
14:
7856:
2520:Holgado, B.; Pêgas, R.V. (2020).
1258:
1247:A rostrum fragment diagnostic to
848:to be monotypic, containing only
642:for a snout found in 1992 in the
7807:
7795:
7673:
7667:
7660:
6443:
6435:
6427:
5235:
5229:
4617:
4612:
4086:
4079:
4071:
884:by Holgado and Pêgas moved both
756:(formerly assigned to the genus
141:
41:
4635:
3219:
3082:
2985:
2962:
2714:Martill, David M. (June 2015).
2621:
2360:10.1590/S0001-37652013000100009
2306:Davies, Ella (4 October 2011).
2203:
1179:Also like its close relatives,
1077:(Kellner & Tomida, 2000) =
968:(Kellner & Tomida, 2000) =
2546:
2299:
2231:Timeline of pterosaur research
2208:Similar to most anhanguerids,
1120:
1063:(Mader & Kellner, 1999) =
1:
7845:Fossil taxa described in 1874
2948:10.1080/08912963.2019.1690482
2701:10.1016/j.cretres.2011.09.003
2526:Acta Palaeontologica Polonica
2494:10.1016/j.cretres.2018.10.018
2236:
1271:(now generally considered as
772:, based on its similarity to
2735:10.1016/j.pgeola.2015.03.004
1743:Holgado & Pêgas (2020).
1176:were robust and box-shaped.
914:List of species and synonyms
583:as species within the genus
541:, rejecting the creation by
7:
2219:
1283:. Later, in 2019, Pentland
1265:Coloborhynchus clavirostris
1147:Coloborhynchus clavirostris
1024:Santanadactylus araripensis
974:Kellner & Tomida, 2000
607:Coloborhynchus clavirostris
553:Coloborhynchus clavirostris
257:Coloborhynchus clavirostris
10:
7861:
7840:Taxa named by Richard Owen
3019:10.1038/s41467-020-19022-2
2932:Targaryendraco wiedenrothi
2898:10.1038/s41598-019-54042-z
2860:10.1038/s41598-019-41280-4
2816:10.1038/s41598-019-49789-4
2601:Veldmeijer, A. J. (2002).
2572:10.1007/s12542-022-00642-6
1083:Kellner & Tomida, 2000
632:This changed in 1994 when
503:History and classification
7701:
7657:
7559:
7509:
7441:
7432:
7404:
7380:
7369:
7325:
7290:
7280:
7176:
7114:
7105:
7101:
7090:
7055:
7014:
6979:
6943:
6934:
6900:
6863:
6854:
6827:
6823:
6800:
6744:
6715:
6711:
6700:
6642:
6613:
6604:
6581:
6572:
6513:
6504:
6489:
6485:
6470:
6457:
6424:
6390:
6274:
6232:
6142:
6133:
6095:
6033:
6024:
6009:
6000:
5939:
5901:
5890:
5839:
5802:
5793:
5746:
5737:
5665:
5650:
5641:
5613:
5604:
5513:
5490:
5479:
5468:
5416:
5405:
5370:
5316:
5303:
5292:
5276:
5261:
5248:
5226:
5196:
5180:
5167:
5106:
5091:
5082:
5033:
5005:
4912:
4901:
4857:
4834:
4825:
4773:
4769:
4758:
4736:
4725:
4677:
4662:
4658:
4643:
4630:
4610:
4568:
4495:
4486:
4432:
4382:
4367:
4358:
4347:
4324:
4313:
4251:
4236:
4227:
4205:
4196:
4166:
4162:
4151:
4127:
4112:
4099:
4068:
4018:
3968:
3958:
3910:
3860:
3851:
3794:
3785:
3746:
3737:
3726:
3691:
3680:
3664:
3652:
3589:
3574:
3570:
3559:
3543:
3531:
3491:
3475:
3466:
3424:
3415:
3383:
3372:
3329:
3298:
3288:
3261:
3246:
3242:
3227:
3214:
3182:
3164:
3151:
3090:
2979:10.1130/abs/2017AM-305496
2762:10.1017/S1755691013000303
2605:Coloborhynchus spielbergi
2449:10.1080/00222931408693521
2257:"Taxonomic review of the
2150:
2124:
2107:
2100:
2083:
2076:
2050:
2033:
2016:
2009:
2002:
1955:
1938:
1931:
1914:
1907:
1890:
1883:
1845:
1828:
1821:
1795:
1778:
1771:
1764:
1754:
1639:
1622:
1615:
1598:
1591:
1574:
1567:
1547:
1540:
1511:
1494:
1487:
1477:
1457:
1450:
1424:
1407:
1400:
1393:
1386:
1369:
1362:
1186:Siroccopteryx moroccensis
1129:1914 illustration of the
1066:Siroccopteryx moroccensis
780:). Kellner, who regarded
765:Siroccopteryx moroccensis
740:Coloborhynchus spielbergi
700:in 2001 by Fastnacht, as
323:
316:
272:
267:
251:
244:
138:Scientific classification
136:
124:
115:
23:
2274:10.3897/zookeys.308.5559
2226:List of pterosaur genera
1309:Uktenadactylus wadleighi
1210:of the possible species
1069:Mader & Kellner 1999
1010:Pterodactylus sedgwickii
950:(Dalla Vecchia, 1993) =
639:Coloborhynchus wadleighi
453:belonging to the family
4650:Archaeopterodactyloidea
2971:Abstracts with Programs
1291:is the sister taxon of
860:named a new species of
702:Coloborhynchus robustus
625:as invalid relative to
414:Nicorhynchus fluviferox
2539:10.4202/app.00751.2020
2427:Hooley, R. W. (1914).
1334:
1251:sp. is known from the
1214:
1134:
1043:Tropeognathus robustus
926:
681:Tropeognathus robustus
593:Reginald Walter Hooley
577:Ornithocheirus cuvieri
519:
7768:Paleobiology Database
2998:Nature Communications
2685:Coloborhynchus capito
1329:
1219:Coloborhynchus capito
1217:The possible species
1206:
1128:
1041:(Wellnhofer, 1987) =
1022:(Wellnhofer, 1985) =
991:Pterodactylus cuvieri
921:
748:Anhanguera spielbergi
510:
4262:Archaeoistiodactylus
4172:Archaeoistiodactylus
1277:Ornithocheirus simus
982:(Bowerbank, 1851) =
957:Dalla Vecchia, 1993
731:as valid species of
7327:Targaryendraconidae
7016:Ornithocheiriformes
6703:Ornithocheiromorpha
6002:Neopterodactyloidea
5929:Dsungaripteromorpha
3697:Campylognathoididae
3417:Eudimorphodontoidea
3010:2020NatCo..11.5293B
2889:2019NatSR...917875K
2808:2019NatSR...913454P
2689:Cretaceous Research
2482:Cretaceous Research
1229:Cambridge Greensand
1080:Anhanguera piscator
971:Anhanguera piscator
870:Ifezouane Formation
797:Cambridge Greensand
729:Anhanguera piscator
725:Anhanguera robustus
543:Harry Govier Seeley
6856:Istiodactyliformes
5330:Lonchognathosaurus
5173:Eupterodactyloidea
4316:Pterodactyliformes
4119:Pterodactylomorpha
4106:Pterodactylomorpha
4020:Pterodactylomorpha
3862:Angustinaripterini
3493:Eudimorphodontinae
3468:Eudimorphodontidae
2936:Historical Biology
2877:Scientific Reports
2849:Scientific Reports
2796:Scientific Reports
2790:Ferrodraco lentoni
2651:10.7717/peerj.3285
2471:"A new species of
2318:on 6 October 2011.
1335:
1319:within the family
1215:
1135:
927:
856:. In 2018, Jacobs
611:Criorhynchus simus
598:Criorhynchus simus
551:, named a species
520:
465:is known from the
132:in multiple views
7783:
7782:
7695:Taxon identifiers
7686:
7685:
7682:
7681:
7655:
7654:
7651:
7650:
7647:
7646:
7643:
7642:
7639:
7638:
7635:
7634:
7631:
7630:
7627:
7626:
7623:
7622:
7365:
7364:
7361:
7360:
7282:Targaryendraconia
7276:
7275:
7086:
7085:
7010:
7009:
7006:
7005:
6796:
6795:
6696:
6695:
6692:
6691:
6688:
6687:
6684:
6683:
6453:
6452:
6422:
6421:
6418:
6417:
6414:
6413:
6386:
6385:
6382:
6381:
6378:
6377:
6374:
6373:
6370:
6369:
6366:
6365:
6362:
6361:
6358:
6357:
6129:
6128:
6097:Chaoyangopterinae
6026:Chaoyangopteridae
5886:
5885:
5882:
5881:
5878:
5877:
5874:
5873:
5870:
5869:
5866:
5865:
5401:
5400:
5397:
5396:
5268:Ornithocheiroidea
5255:Ornithocheiroidea
5244:
5243:
5224:
5223:
5220:
5219:
5216:
5215:
5198:Ornithocheiroidea
5163:
5162:
5159:
5158:
5155:
5154:
5151:
5150:
5147:
5146:
4961:Gladocephaloideus
4897:
4896:
4893:
4892:
4859:Aurorazhdarchidae
4761:Ctenochasmatoidea
4721:
4720:
4679:Germanodactylidae
4626:
4625:
4608:
4607:
4604:
4603:
4600:
4599:
4596:
4595:
4592:
4591:
4588:
4587:
4482:
4481:
4457:Dendrorhynchoides
4434:Batrachognathinae
4400:Dendrorhynchoides
4330:Changchengopterus
4309:
4308:
4305:
4304:
4301:
4300:
4095:
4094:
4066:
4065:
4062:
4061:
4058:
4057:
4054:
4053:
4050:
4049:
4046:
4045:
4042:
4041:
4038:
4037:
3954:
3953:
3950:
3949:
3946:
3945:
3871:Angustinaripterus
3787:Rhamphorhynchinae
3739:Rhamphorhynchidae
3722:
3721:
3713:Campylognathoides
3648:
3647:
3527:
3526:
3523:
3522:
3519:
3518:
3426:Raeticodactylidae
3368:
3367:
3364:
3363:
3331:Austriadraconidae
3210:
3209:
3206:
3205:
3202:
3201:
2200:
2199:
2192:
2191:
2183:
2182:
2174:
2173:
2165:
2164:
2139:
2138:
2065:
2064:
1988:
1987:
1979:
1978:
1970:
1969:
1869:
1868:
1860:
1859:
1810:
1809:
1735:
1734:
1726:
1725:
1717:
1716:
1708:
1707:
1699:
1698:
1690:
1689:
1681:
1680:
1672:
1671:
1663:
1662:
1654:
1653:
1526:
1525:
1439:
1438:
1194:Alexander Kellner
1156:Like the related
1116:
1108:
1099:(Seeley, 1870) =
1093:
1091:Veldmeijer, 2003
1057:
1047:
1045:Wellnhofer, 1987
1032:
1026:Wellnhofer, 1985
1014:
1008:(Newton, 1888) =
995:
989:(Newton, 1888) =
976:
959:
776:(type species of
690:Santana Formation
648:Paw Paw Formation
609:was a synonym of
461:in some studies.
435:
434:
423:
394:
382:
354:
310:
291:
240:
7852:
7812:
7811:
7810:
7800:
7799:
7798:
7791:
7776:
7775:
7763:
7762:
7750:
7749:
7737:
7736:
7735:
7722:
7721:
7720:
7690:
7689:
7677:
7671:
7665:
7664:
7511:Coloborhynchinae
7439:
7438:
7394:Ornithocheiridae
7386:Brasileodactylus
7378:
7377:
7288:
7287:
7178:Ornithocheirinae
7112:
7111:
7107:Ornithocheiridae
7103:
7102:
7099:
7098:
7057:
7056:
7053:
7052:
7032:Brasileodactylus
6941:
6940:
6861:
6860:
6829:Lonchodraconidae
6825:
6824:
6821:
6820:
6713:
6712:
6709:
6708:
6611:
6610:
6579:
6578:
6574:Nyctosauromorpha
6511:
6510:
6487:
6486:
6483:
6482:
6477:Pteranodontoidea
6472:
6471:
6464:Pteranodontoidea
6459:
6458:
6448:
6447:
6440:
6439:
6432:
6431:
6396:Pteranodontoidea
6392:
6391:
6276:Quetzalcoatlinae
6265:Mistralazhdarcho
6230:
6229:
6171:
6170:
6140:
6139:
6031:
6030:
6007:
6006:
5899:
5898:
5800:
5799:
5744:
5743:
5663:
5662:
5648:
5647:
5611:
5610:
5515:Thalassodromidae
5488:
5487:
5477:
5476:
5414:
5413:
5372:Dsungaripterinae
5314:
5313:
5309:Dsungaripteridae
5305:
5304:
5301:
5300:
5274:
5273:
5263:
5262:
5250:
5249:
5239:
5234:
5233:
5178:
5177:
5169:
5168:
5089:
5088:
5084:Ctenochasmatinae
5066:Plataleorhynchus
4910:
4909:
4904:Ctenochasmatidae
4832:
4831:
4771:
4770:
4767:
4766:
4734:
4733:
4728:Euctenochasmatia
4660:
4659:
4656:
4655:
4645:
4644:
4632:
4631:
4621:
4616:
4544:Propterodactylus
4530:Ningchengopterus
4493:
4492:
4488:Pterodactyloidea
4365:
4364:
4356:
4355:
4322:
4321:
4234:
4233:
4203:
4202:
4164:
4163:
4160:
4159:
4125:
4124:
4114:
4113:
4101:
4100:
4090:
4084:
4083:
4076:
4075:
3994:Jianchangopterus
3980:Fenghuangopterus
3966:
3965:
3912:Rhamphorhynchini
3858:
3857:
3792:
3791:
3744:
3743:
3735:
3734:
3693:
3692:
3689:
3688:
3662:
3661:
3591:Dimorphodontidae
3572:
3571:
3568:
3567:
3541:
3540:
3473:
3472:
3422:
3421:
3381:
3380:
3296:
3295:
3244:
3243:
3240:
3239:
3229:
3228:
3216:
3215:
3162:
3161:
3153:
3152:
3146:
3145:
3141:Pterosauromorpha
3092:
3091:
3077:
3070:
3063:
3054:
3053:
3042:
3041:
3031:
3021:
2989:
2983:
2982:
2966:
2960:
2959:
2942:(8): 1266–1280.
2927:
2921:
2920:
2910:
2900:
2868:
2862:
2844:
2838:
2837:
2827:
2783:
2774:
2773:
2756:(3–4): 383–398.
2745:
2739:
2738:
2720:
2711:
2705:
2704:
2680:
2674:
2673:
2663:
2653:
2625:
2619:
2618:
2603:"Description of
2598:
2585:
2584:
2574:
2550:
2544:
2543:
2541:
2517:
2506:
2505:
2479:
2466:
2453:
2452:
2424:
2418:
2417:
2407:
2394:
2373:
2372:
2362:
2329:
2320:
2319:
2314:. Archived from
2303:
2297:
2296:
2286:
2276:
2252:
2103:
2102:
2079:
2078:
2012:
2011:
2005:
2004:
1934:
1933:
1910:
1909:
1886:
1885:
1880:Coloborhynchinae
1824:
1823:
1774:
1773:
1767:
1766:
1757:
1756:
1747:
1746:
1618:
1617:
1594:
1593:
1570:
1569:
1564:Ornithocheirinae
1543:
1542:
1537:Ornithocheiridae
1490:
1489:
1480:
1479:
1453:
1452:
1411:Brasileodactylus
1403:
1402:
1396:
1395:
1389:
1388:
1365:
1364:
1353:
1352:
1338:
1253:Wessex Formation
1208:Life restoration
1115:
1107:
1092:
1056:
1046:
1027:
1013:
994:
993:Bowerbank, 1851
975:
958:
940:Owen, 1874, the
892:to a new genus,
882:Coloborhynchinae
833:compressirostris
696:was assigned to
467:Lower Cretaceous
459:Ornithocheiridae
417:
392:
380:
346:
327:List of synonyms
304:
296:
285:
277:
254:
235:
228:
215:
201:Pterodactyloidea
199:
186:
146:
145:
128:jaw fragment of
120:
110:
40:
31:Early Cretaceous
29:Temporal range:
21:
20:
7860:
7859:
7855:
7854:
7853:
7851:
7850:
7849:
7830:Pteranodontoids
7820:
7819:
7818:
7808:
7806:
7796:
7794:
7786:
7784:
7779:
7771:
7766:
7758:
7753:
7745:
7740:
7731:
7730:
7725:
7716:
7715:
7710:
7697:
7687:
7678:
7659:
7619:
7600:Liaoningopterus
7555:
7505:
7443:Tropeognathinae
7428:
7400:
7374:
7357:
7321:
7292:Cimoliopteridae
7272:
7172:
7095:
7082:
7077:Zhenyuanopterus
7051:
7002:
6981:Istiodactylinae
6975:
6956:Longchengpterus
6949:Lingyuanopterus
6936:Istiodactylidae
6930:
6896:
6869:Hongshanopterus
6850:
6819:
6805:
6792:
6755:Hongshanopterus
6740:
6705:
6680:
6653:Barbaridactylus
6638:
6600:
6568:
6515:Pteranodontidae
6500:
6495:Santanadactylus
6479:
6466:
6449:
6442:
6434:
6426:
6410:
6354:
6349:Zhejiangopterus
6270:
6228:
6207:Palaeocursornis
6169:
6148:Montanazhdarcho
6135:Azhdarchiformes
6125:
6106:Chaoyangopterus
6091:
6020:
5996:
5982:Montanazhdarcho
5935:
5895:
5893:Azhdarchomorpha
5862:
5835:
5789:
5752:Aymberedactylus
5733:
5661:
5637:
5625:Aymberedactylus
5600:
5580:Thalassodromeus
5509:
5484:
5473:
5464:
5452:Montanazhdarcho
5410:
5393:
5381:Domeykodactylus
5366:
5360:Tendaguripterus
5297:
5288:
5270:
5257:
5240:
5228:
5212:
5192:
5186:Altmuehlopterus
5143:
5102:
5078:
5029:
5001:
4906:
4889:
4853:
4847:Gallodactylidae
4827:Aurorazhdarchia
4821:
4799:Normannognathus
4775:Gallodactylidae
4763:
4754:
4730:
4717:
4711:Tendaguripterus
4703:Normannognathus
4696:Germanodactylus
4688:Altmuehlopterus
4673:
4652:
4639:
4622:
4584:
4564:
4478:
4443:Batrachognathus
4428:
4423:Vesperopterylus
4378:
4352:
4343:
4318:
4297:
4253:Wukongopterinae
4247:
4229:Wukongopteridae
4223:
4192:
4179:Normannognathus
4156:
4147:
4121:
4108:
4091:
4078:
4070:
4034:
4014:
3987:Jianchangnathus
3970:Scaphognathidae
3960:Digibrevisauria
3942:
3937:Rhamphorhynchus
3906:
3893:Qinglongopterus
3885:Harpactognathus
3853:Rhamphorhynchae
3847:
3842:Qinglongopterus
3821:Harpactognathus
3781:
3775:Scaphognathinae
3767:Parapsicephalus
3731:
3718:
3706:Bergamodactylus
3685:
3676:
3658:
3644:
3622:Parapsicephalus
3585:
3564:
3562:Macronychoptera
3555:
3537:
3515:
3487:
3481:Arcticodactylus
3462:
3450:Raeticodactylus
3411:
3377:
3360:
3340:Arcticodactylus
3325:
3319:Raeticodactylus
3284:
3272:Austriadactylus
3257:
3236:
3223:
3198:
3178:
3172:Avemetatarsalia
3157:Avemetatarsalia
3147:
3132:Avemetatarsalia
3097:
3096:
3086:
3081:
3046:
3045:
2990:
2986:
2967:
2963:
2928:
2924:
2869:
2865:
2845:
2841:
2784:
2777:
2746:
2742:
2718:
2712:
2708:
2681:
2677:
2626:
2622:
2617:(3003): 35–139.
2611:Scripta Geology
2599:
2588:
2551:
2547:
2518:
2509:
2477:
2467:
2456:
2443:(78): 529–557.
2425:
2421:
2405:
2395:
2376:
2330:
2323:
2304:
2300:
2253:
2244:
2239:
2222:
2206:
2201:
2193:
2184:
2175:
2166:
2140:
2087:Liaoningopterus
2066:
1989:
1980:
1971:
1870:
1861:
1811:
1761:Tropeognathinae
1736:
1727:
1718:
1709:
1700:
1691:
1682:
1673:
1664:
1655:
1527:
1498:Liaoningopterus
1440:
1331:C. clavirostris
1261:
1198:C. clavirostris
1131:C. clavirostris
1123:
938:C. clavirostris
916:
850:C. clavirostris
825:C. clavirostris
817:C. clavirostris
759:Santanadactylus
744:C. clavirostris
721:C. clavirostris
686:Romualdo Member
667:C. clavirostris
615:C. clavirostris
524:ornithocheiroid
516:C. clavirostris
505:
431:
428:
416:
408:
407:
399:
391:
379:
368:
367:
359:
345:
343:"Ptenodactylus"
337:
336:
329:
328:
303:
284:
263:
260:
234:
226:
213:
197:
184:
140:
130:C. clavirostris
111:
109:
108:
107:
106:
101:
96:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
35:
34:
27:
17:
12:
11:
5:
7858:
7848:
7847:
7842:
7837:
7832:
7817:
7816:
7814:United Kingdom
7804:
7781:
7780:
7778:
7777:
7764:
7751:
7738:
7733:Coloborhynchus
7723:
7707:
7705:
7703:Coloborhynchus
7699:
7698:
7684:
7683:
7680:
7679:
7658:
7656:
7653:
7652:
7649:
7648:
7645:
7644:
7641:
7640:
7637:
7636:
7633:
7632:
7629:
7628:
7625:
7624:
7621:
7620:
7618:
7617:
7610:
7603:
7596:
7589:
7581:
7577:Caulkicephalus
7573:
7565:
7563:
7557:
7556:
7554:
7553:
7550:Uktenadactylus
7546:
7538:
7531:
7528:Coloborhynchus
7524:
7516:
7514:
7507:
7506:
7504:
7503:
7496:
7489:
7481:
7473:
7465:
7457:
7448:
7446:
7436:
7430:
7429:
7427:
7426:
7419:
7411:
7409:
7402:
7401:
7399:
7398:
7390:
7381:
7375:
7370:
7367:
7366:
7363:
7362:
7359:
7358:
7356:
7355:
7352:Targaryendraco
7348:
7340:
7331:
7329:
7323:
7322:
7320:
7319:
7312:
7304:
7296:
7294:
7285:
7278:
7277:
7274:
7273:
7271:
7270:
7266:Uktenadactylus
7262:
7254:
7251:Ornithocheirus
7247:
7239:
7231:
7223:
7215:
7207:
7203:Coloborhynchus
7199:
7191:
7182:
7180:
7174:
7173:
7171:
7170:
7162:
7154:
7146:
7142:Caulkicephalus
7138:
7130:
7127:Arthurdactylus
7123:
7115:
7109:
7096:
7093:Ornithocheirae
7091:
7088:
7087:
7084:
7083:
7081:
7080:
7073:
7065:
7063:
7050:
7049:
7042:
7035:
7028:
7020:
7018:
7012:
7011:
7008:
7007:
7004:
7003:
7001:
7000:
6993:
6985:
6983:
6977:
6976:
6974:
6973:
6966:
6959:
6952:
6944:
6938:
6932:
6931:
6929:
6928:
6921:
6918:Linlongopterus
6914:
6906:
6904:
6902:Mimodactylidae
6898:
6897:
6895:
6894:
6890:Lonchodectidae
6886:
6879:
6876:Linlongopterus
6872:
6864:
6858:
6852:
6851:
6849:
6848:
6841:
6833:
6831:
6818:
6817:
6808:
6806:
6801:
6798:
6797:
6794:
6793:
6791:
6790:
6786:Targaryendraco
6782:
6774:
6767:
6759:
6750:
6748:
6746:Lonchodectidae
6742:
6741:
6739:
6738:
6731:
6724:
6716:
6706:
6701:
6698:
6697:
6694:
6693:
6690:
6689:
6686:
6685:
6682:
6681:
6679:
6678:
6670:
6663:
6660:Muzquizopteryx
6656:
6648:
6646:
6640:
6639:
6637:
6636:
6629:
6622:
6614:
6608:
6606:Aponyctosauria
6602:
6601:
6599:
6598:
6590:
6582:
6576:
6570:
6569:
6567:
6566:
6558:
6550:
6543:
6535:
6528:
6519:
6517:
6508:
6502:
6501:
6499:
6498:
6490:
6480:
6475:
6468:
6467:
6455:
6454:
6451:
6450:
6425:
6423:
6420:
6419:
6416:
6415:
6412:
6411:
6409:
6408:
6407:
6406:
6400:
6398:
6388:
6387:
6384:
6383:
6380:
6379:
6376:
6375:
6372:
6371:
6368:
6367:
6364:
6363:
6360:
6359:
6356:
6355:
6353:
6352:
6345:
6337:
6334:Thanatosdrakon
6330:
6327:Quetzalcoatlus
6323:
6320:Phosphatodraco
6316:
6309:
6302:
6295:
6292:Arambourgiania
6288:
6280:
6278:
6272:
6271:
6269:
6268:
6261:
6254:
6247:
6238:
6236:
6227:
6226:
6218:
6210:
6203:
6199:Navajodactylus
6195:
6188:
6179:
6177:
6168:
6167:
6159:
6152:
6143:
6137:
6131:
6130:
6127:
6126:
6124:
6123:
6120:Shenzhoupterus
6116:
6109:
6101:
6099:
6093:
6092:
6090:
6089:
6081:
6073:
6066:
6059:
6051:
6047:Argentinadraco
6043:
6034:
6028:
6022:
6021:
6019:
6018:
6010:
6004:
5998:
5997:
5995:
5994:
5986:
5978:
5970:
5962:
5958:Argentinadraco
5954:
5946:
5944:
5937:
5936:
5934:
5933:
5925:
5918:
5911:
5902:
5896:
5891:
5888:
5887:
5884:
5883:
5880:
5879:
5876:
5875:
5872:
5871:
5868:
5867:
5864:
5863:
5861:
5860:
5853:
5845:
5843:
5837:
5836:
5834:
5833:
5826:
5819:
5812:
5803:
5797:
5791:
5790:
5788:
5787:
5780:
5772:
5764:
5756:
5747:
5741:
5735:
5734:
5732:
5731:
5724:
5717:
5714:Nemicolopterus
5710:
5703:
5696:
5688:
5680:
5671:
5669:
5660:
5659:
5651:
5645:
5639:
5638:
5636:
5635:
5628:
5620:
5618:
5608:
5606:Tapejariformes
5602:
5601:
5599:
5598:
5590:
5583:
5576:
5568:
5560:
5553:
5545:
5541:Argentinadraco
5537:
5529:
5520:
5518:
5511:
5510:
5508:
5507:
5499:
5491:
5485:
5482:Tapejaromorpha
5480:
5474:
5469:
5466:
5465:
5463:
5462:
5455:
5448:
5440:
5432:
5425:
5422:Argentinadraco
5417:
5411:
5406:
5403:
5402:
5399:
5398:
5395:
5394:
5392:
5391:
5388:Dsungaripterus
5384:
5376:
5374:
5368:
5367:
5365:
5364:
5356:
5348:
5341:
5334:
5326:
5317:
5311:
5298:
5293:
5290:
5289:
5287:
5286:
5277:
5271:
5266:
5259:
5258:
5246:
5245:
5242:
5241:
5227:
5225:
5222:
5221:
5218:
5217:
5214:
5213:
5211:
5210:
5209:
5208:
5202:
5200:
5194:
5193:
5191:
5190:
5181:
5175:
5165:
5164:
5161:
5160:
5157:
5156:
5153:
5152:
5149:
5148:
5145:
5144:
5142:
5141:
5134:
5127:
5120:
5112:
5110:
5108:Pterodaustrini
5104:
5103:
5101:
5100:
5092:
5086:
5080:
5079:
5077:
5076:
5069:
5062:
5055:
5047:
5039:
5037:
5035:Gnathosaurinae
5031:
5030:
5028:
5027:
5020:
5012:
5010:
5007:Moganopterinae
5003:
5002:
5000:
4999:
4992:
4985:
4978:
4971:
4964:
4957:
4950:
4943:
4936:
4929:
4926:Balaenognathus
4922:
4913:
4907:
4902:
4899:
4898:
4895:
4894:
4891:
4890:
4888:
4887:
4879:
4876:Aurorazhdarcho
4872:
4863:
4861:
4855:
4854:
4852:
4851:
4843:
4835:
4829:
4823:
4822:
4820:
4819:
4815:Moganopterinae
4811:
4803:
4795:
4788:
4784:Aurorazhdarcho
4779:
4777:
4764:
4759:
4756:
4755:
4753:
4752:
4745:
4737:
4731:
4726:
4723:
4722:
4719:
4718:
4716:
4715:
4707:
4699:
4692:
4683:
4681:
4675:
4674:
4672:
4671:
4663:
4653:
4648:
4641:
4640:
4628:
4627:
4624:
4623:
4611:
4609:
4606:
4605:
4602:
4601:
4598:
4597:
4594:
4593:
4590:
4589:
4586:
4585:
4583:
4582:
4581:
4580:
4574:
4572:
4566:
4565:
4563:
4562:
4555:
4547:
4540:
4533:
4526:
4519:
4511:
4504:
4496:
4490:
4484:
4483:
4480:
4479:
4477:
4476:
4469:
4461:
4453:
4446:
4438:
4436:
4430:
4429:
4427:
4426:
4419:
4412:
4404:
4396:
4388:
4386:
4384:Anurognathinae
4380:
4379:
4377:
4376:
4368:
4362:
4360:Anurognathidae
4353:
4348:
4345:
4344:
4342:
4341:
4338:Douzhanopterus
4334:
4325:
4319:
4314:
4311:
4310:
4307:
4306:
4303:
4302:
4299:
4298:
4296:
4295:
4288:
4284:Douzhanopterus
4280:
4273:
4266:
4257:
4255:
4249:
4248:
4246:
4245:
4242:Kunpengopterus
4237:
4231:
4225:
4224:
4222:
4221:
4214:
4206:
4200:
4194:
4193:
4191:
4190:
4186:Anurognathidae
4182:
4175:
4167:
4157:
4154:Monofenestrata
4152:
4149:
4148:
4146:
4145:
4137:
4128:
4122:
4117:
4110:
4109:
4097:
4096:
4093:
4092:
4069:
4067:
4064:
4063:
4060:
4059:
4056:
4055:
4052:
4051:
4048:
4047:
4044:
4043:
4040:
4039:
4036:
4035:
4033:
4032:
4031:
4030:
4024:
4022:
4016:
4015:
4013:
4012:
4004:
3997:
3990:
3983:
3975:
3973:
3963:
3956:
3955:
3952:
3951:
3948:
3947:
3944:
3943:
3941:
3940:
3933:
3925:
3916:
3914:
3908:
3907:
3905:
3904:
3897:
3889:
3881:
3874:
3866:
3864:
3855:
3849:
3848:
3846:
3845:
3838:
3835:Orientognathus
3831:
3824:
3817:
3810:
3803:
3795:
3789:
3783:
3782:
3780:
3779:
3771:
3763:
3756:
3752:Dolicorhamphus
3747:
3741:
3732:
3729:Breviquartossa
3727:
3724:
3723:
3720:
3719:
3717:
3716:
3709:
3701:
3699:
3686:
3681:
3678:
3677:
3675:
3674:
3665:
3659:
3653:
3650:
3649:
3646:
3645:
3643:
3642:
3634:
3626:
3618:
3611:
3608:Caelestiventus
3604:
3595:
3593:
3587:
3586:
3584:
3583:
3575:
3565:
3560:
3557:
3556:
3554:
3553:
3544:
3538:
3534:Zambellisauria
3532:
3529:
3528:
3525:
3524:
3521:
3520:
3517:
3516:
3514:
3513:
3506:
3502:Carniadactylus
3497:
3495:
3489:
3488:
3486:
3485:
3476:
3470:
3464:
3463:
3461:
3460:
3453:
3446:
3439:
3430:
3428:
3419:
3413:
3412:
3410:
3409:
3401:
3393:
3384:
3378:
3373:
3370:
3369:
3366:
3365:
3362:
3361:
3359:
3358:
3355:Seazzadactylus
3351:
3344:
3335:
3333:
3327:
3326:
3324:
3323:
3315:
3308:
3304:Carniadactylus
3299:
3293:
3286:
3285:
3283:
3282:
3275:
3267:
3265:
3259:
3258:
3256:
3255:
3252:Daohugoupterus
3247:
3237:
3232:
3225:
3224:
3212:
3211:
3208:
3207:
3204:
3203:
3200:
3199:
3197:
3196:
3195:
3194:
3188:
3186:
3180:
3179:
3177:
3176:
3175:
3174:
3165:
3159:
3149:
3148:
3144:
3143:
3134:
3125:
3116:
3110:
3104:
3088:
3087:
3080:
3079:
3072:
3065:
3057:
3044:
3043:
2984:
2961:
2922:
2863:
2839:
2775:
2740:
2729:(3): 377–380.
2706:
2675:
2620:
2586:
2565:(3): 519–568.
2545:
2507:
2473:Coloborhynchus
2454:
2431:Ornithocheirus
2419:
2401:Coloborhynchus
2374:
2353:(1): 113–135.
2341:T. mesembrinus
2321:
2298:
2267:(308): 1–112.
2259:Ornithocheirus
2241:
2240:
2238:
2235:
2234:
2233:
2228:
2221:
2218:
2210:Coloborhynchus
2205:
2202:
2198:
2197:
2190:
2189:
2186:
2185:
2181:
2180:
2177:
2176:
2172:
2171:
2168:
2167:
2163:
2162:
2159:
2158:
2149:
2146:
2145:
2142:
2141:
2137:
2136:
2133:
2132:
2123:
2120:
2119:
2116:
2115:
2106:
2101:
2099:
2096:
2095:
2092:
2091:
2082:
2077:
2075:
2072:
2071:
2068:
2067:
2063:
2062:
2059:
2058:
2049:
2046:
2045:
2042:
2041:
2032:
2029:
2028:
2025:
2024:
2020:Caulkicephalus
2015:
2010:
2008:
2003:
2001:
1995:
1994:
1991:
1990:
1986:
1985:
1982:
1981:
1977:
1976:
1973:
1972:
1968:
1967:
1964:
1963:
1959:Uktenadactylus
1954:
1951:
1950:
1947:
1946:
1937:
1932:
1930:
1927:
1926:
1923:
1922:
1918:Coloborhynchus
1913:
1908:
1906:
1903:
1902:
1899:
1898:
1889:
1884:
1882:
1876:
1875:
1872:
1871:
1867:
1866:
1863:
1862:
1858:
1857:
1854:
1853:
1844:
1841:
1840:
1837:
1836:
1827:
1822:
1820:
1817:
1816:
1813:
1812:
1808:
1807:
1804:
1803:
1794:
1791:
1790:
1787:
1786:
1777:
1772:
1770:
1765:
1763:
1755:
1753:
1745:
1737:
1733:
1732:
1729:
1728:
1724:
1723:
1720:
1719:
1715:
1714:
1711:
1710:
1706:
1705:
1702:
1701:
1697:
1696:
1693:
1692:
1688:
1687:
1684:
1683:
1679:
1678:
1675:
1674:
1670:
1669:
1666:
1665:
1661:
1660:
1657:
1656:
1652:
1651:
1648:
1647:
1638:
1635:
1634:
1631:
1630:
1621:
1616:
1614:
1611:
1610:
1607:
1606:
1602:Ornithocheirus
1597:
1592:
1590:
1587:
1586:
1583:
1582:
1578:Coloborhynchus
1573:
1568:
1566:
1560:
1559:
1556:
1555:
1546:
1541:
1539:
1533:
1532:
1529:
1528:
1524:
1523:
1520:
1519:
1510:
1507:
1506:
1503:
1502:
1493:
1488:
1486:
1478:
1476:
1474:Ornithocheirae
1470:
1469:
1466:
1465:
1456:
1451:
1449:
1446:
1445:
1442:
1441:
1437:
1436:
1433:
1432:
1423:
1420:
1419:
1416:
1415:
1406:
1401:
1399:
1394:
1392:
1387:
1385:
1382:
1381:
1378:
1377:
1368:
1363:
1361:
1351:
1336:
1317:Coloborhynchus
1301:Coloborhynchus
1293:Ornithocheirus
1289:Coloborhynchus
1281:Ornithocheirae
1273:Uktenadactylus
1260:
1259:Classification
1257:
1249:Coloborhynchus
1242:Coloborhynchus
1181:Coloborhynchus
1174:Coloborhynchus
1165:Uktenadactylus
1151:Coloborhynchus
1143:Coloborhynchus
1122:
1119:
1118:
1117:
1109:
1102:Ornithocheirus
1094:
1084:
1070:
1061:C. moroccensis
1058:
1048:
1033:
1028:now seen as a
1019:C. araripensis
1015:
1004:Ornithocheirus
1001:(Owen 1859) =
996:
985:Ornithocheirus
977:
960:
944:
931:Coloborhynchus
915:
912:
908:Coloborhynchus
862:Coloborhynchus
846:Coloborhynchus
829:Ornithocheirus
801:Coloborhynchus
790:Coloborhynchus
782:Uktenadactylus
778:Uktenadactylus
770:Coloborhynchus
753:C. araripensis
698:Coloborhynchus
672:Uktenadactylus
659:Coloborhynchus
655:Coloborhynchus
623:Coloborhynchus
619:Ornithocheirus
603:Ornithocheirus
585:Coloborhycnhus
548:Ornithocheirus
532:Coloborhynchus
528:Coloborhynchus
504:
501:
487:Coloborhynchus
463:Coloborhynchus
448:pterodactyloid
439:Coloborhynchus
433:
432:
430:
429:
427:
426:
425:
424:
402:
401:
400:
398:
397:
396:
395:
393:(Seeley, 1870)
383:
375:Ornithocheirus
362:
361:
360:
358:
357:
356:
355:
347:Seeley, 1869 (
335:Genus synonymy
334:
333:
332:
326:
325:
324:
321:
320:
314:
313:
312:
311:
292:
270:
269:
268:Other species
265:
264:
261:
249:
248:
242:
241:
231:Coloborhynchus
224:
220:
219:
217:Ornithocheirae
211:
204:
203:
195:
191:
190:
182:
178:
177:
172:
168:
167:
162:
158:
157:
152:
148:
147:
134:
133:
122:
121:
113:
112:
104:
103:
102:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
42:
28:
25:Coloborhynchus
15:
9:
6:
4:
3:
2:
7857:
7846:
7843:
7841:
7838:
7836:
7833:
7831:
7828:
7827:
7825:
7815:
7805:
7803:
7793:
7792:
7789:
7774:
7769:
7765:
7761:
7756:
7752:
7748:
7743:
7739:
7734:
7728:
7724:
7719:
7713:
7709:
7708:
7706:
7704:
7700:
7696:
7691:
7676:
7672:
7670:
7663:
7616:
7615:
7614:Maaradactylus
7611:
7609:
7608:
7604:
7602:
7601:
7597:
7595:
7594:
7590:
7587:
7586:
7585:Cearadactylus
7582:
7579:
7578:
7574:
7572:
7571:
7567:
7566:
7564:
7562:
7561:Anhanguerinae
7558:
7552:
7551:
7547:
7544:
7543:
7542:Siroccopteryx
7539:
7537:
7536:
7532:
7530:
7529:
7525:
7523:
7522:
7518:
7517:
7515:
7512:
7508:
7502:
7501:
7500:Tropeognathus
7497:
7495:
7494:
7490:
7487:
7486:
7485:Siroccopteryx
7482:
7479:
7478:
7474:
7471:
7470:
7466:
7463:
7462:
7458:
7455:
7454:
7450:
7449:
7447:
7444:
7440:
7437:
7435:
7434:Anhangueridae
7431:
7425:
7424:
7423:Iberodactylus
7420:
7418:
7417:
7413:
7412:
7410:
7407:
7403:
7396:
7395:
7391:
7388:
7387:
7383:
7382:
7379:
7376:
7373:
7368:
7354:
7353:
7349:
7346:
7345:
7341:
7338:
7337:
7333:
7332:
7330:
7328:
7324:
7318:
7317:
7316:Cimoliopterus
7313:
7310:
7309:
7308:Camposipterus
7305:
7303:
7302:
7298:
7297:
7295:
7293:
7289:
7286:
7283:
7279:
7268:
7267:
7263:
7260:
7259:
7258:Siroccopteryx
7255:
7253:
7252:
7248:
7245:
7244:
7240:
7237:
7236:
7232:
7229:
7228:
7224:
7221:
7220:
7216:
7213:
7212:
7208:
7205:
7204:
7200:
7197:
7196:
7195:Camposipterus
7192:
7189:
7188:
7184:
7183:
7181:
7179:
7175:
7168:
7167:
7166:Tropeognathus
7163:
7160:
7159:
7155:
7152:
7151:
7150:Cimoliopterus
7147:
7144:
7143:
7139:
7136:
7135:
7134:Camposipterus
7131:
7129:
7128:
7124:
7122:
7121:
7120:Araripesaurus
7117:
7116:
7113:
7110:
7108:
7104:
7100:
7097:
7094:
7089:
7079:
7078:
7074:
7072:
7071:
7067:
7066:
7064:
7062:
7061:Boreopteridae
7058:
7054:
7048:
7047:
7043:
7041:
7040:
7039:Cearadactylus
7036:
7034:
7033:
7029:
7027:
7026:
7022:
7021:
7019:
7017:
7013:
6999:
6998:
6994:
6992:
6991:
6990:Istiodactylus
6987:
6986:
6984:
6982:
6978:
6972:
6971:
6967:
6965:
6964:
6960:
6958:
6957:
6953:
6951:
6950:
6946:
6945:
6942:
6939:
6937:
6933:
6927:
6926:
6922:
6920:
6919:
6915:
6913:
6912:
6908:
6907:
6905:
6903:
6899:
6892:
6891:
6887:
6885:
6884:
6883:Yixianopterus
6880:
6878:
6877:
6873:
6871:
6870:
6866:
6865:
6862:
6859:
6857:
6853:
6847:
6846:
6842:
6840:
6839:
6835:
6834:
6832:
6830:
6826:
6822:
6815:
6814:
6810:
6809:
6807:
6804:
6799:
6788:
6787:
6783:
6780:
6779:
6775:
6773:
6772:
6768:
6765:
6764:
6760:
6757:
6756:
6752:
6751:
6749:
6747:
6743:
6737:
6736:
6732:
6730:
6729:
6725:
6723:
6722:
6718:
6717:
6714:
6710:
6707:
6704:
6699:
6676:
6675:
6671:
6669:
6668:
6664:
6662:
6661:
6657:
6655:
6654:
6650:
6649:
6647:
6645:
6644:Nyctosauridae
6641:
6635:
6634:
6630:
6628:
6627:
6623:
6621:
6620:
6616:
6615:
6612:
6609:
6607:
6603:
6596:
6595:
6591:
6589:
6588:
6587:Alamodactylus
6584:
6583:
6580:
6577:
6575:
6571:
6564:
6563:
6559:
6556:
6555:
6551:
6549:
6548:
6544:
6541:
6540:
6536:
6534:
6533:
6529:
6526:
6525:
6521:
6520:
6518:
6516:
6512:
6509:
6507:
6506:Pteranodontia
6503:
6497:
6496:
6492:
6491:
6488:
6484:
6481:
6478:
6473:
6469:
6465:
6460:
6456:
6446:
6441:
6438:
6430:
6404:
6403:
6402:
6401:
6399:
6397:
6393:
6389:
6351:
6350:
6346:
6343:
6342:
6341:Wellnhopterus
6338:
6336:
6335:
6331:
6329:
6328:
6324:
6322:
6321:
6317:
6315:
6314:
6313:Hatzegopteryx
6310:
6308:
6307:
6303:
6301:
6300:
6296:
6294:
6293:
6289:
6287:
6286:
6285:Aralazhdarcho
6282:
6281:
6279:
6277:
6273:
6267:
6266:
6262:
6260:
6259:
6255:
6253:
6252:
6248:
6245:
6244:
6240:
6239:
6237:
6235:
6231:
6224:
6223:
6219:
6216:
6215:
6211:
6209:
6208:
6204:
6201:
6200:
6196:
6194:
6193:
6189:
6186:
6185:
6181:
6180:
6178:
6176:
6172:
6165:
6164:
6160:
6158:
6157:
6156:Radiodactylus
6153:
6150:
6149:
6145:
6144:
6141:
6138:
6136:
6132:
6122:
6121:
6117:
6115:
6114:
6110:
6108:
6107:
6103:
6102:
6100:
6098:
6094:
6087:
6086:
6082:
6079:
6078:
6074:
6072:
6071:
6067:
6065:
6064:
6060:
6057:
6056:
6052:
6049:
6048:
6044:
6041:
6040:
6039:Apatorhamphus
6036:
6035:
6032:
6029:
6027:
6023:
6017:
6016:
6012:
6011:
6008:
6005:
6003:
5999:
5992:
5991:
5987:
5984:
5983:
5979:
5976:
5975:
5971:
5968:
5967:
5963:
5960:
5959:
5955:
5953:
5952:
5948:
5947:
5945:
5942:
5938:
5931:
5930:
5926:
5924:
5923:
5919:
5917:
5916:
5912:
5909:
5908:
5904:
5903:
5900:
5897:
5894:
5889:
5859:
5858:
5854:
5852:
5851:
5847:
5846:
5844:
5842:
5838:
5832:
5831:
5830:Tupandactylus
5827:
5825:
5824:
5820:
5818:
5817:
5813:
5810:
5809:
5805:
5804:
5801:
5798:
5796:
5792:
5786:
5785:
5781:
5778:
5777:
5773:
5770:
5769:
5765:
5762:
5761:
5760:Caupedactylus
5757:
5754:
5753:
5749:
5748:
5745:
5742:
5740:
5736:
5730:
5729:
5725:
5723:
5722:
5718:
5716:
5715:
5711:
5709:
5708:
5704:
5702:
5701:
5697:
5694:
5693:
5689:
5686:
5685:
5681:
5678:
5677:
5673:
5672:
5670:
5668:
5664:
5658:
5657:
5653:
5652:
5649:
5646:
5644:
5640:
5634:
5633:
5632:Caupedactylus
5629:
5627:
5626:
5622:
5621:
5619:
5616:
5615:Caupedactylia
5612:
5609:
5607:
5603:
5596:
5595:
5591:
5589:
5588:
5584:
5582:
5581:
5577:
5574:
5573:
5569:
5566:
5565:
5561:
5559:
5558:
5554:
5551:
5550:
5546:
5543:
5542:
5538:
5535:
5534:
5530:
5527:
5526:
5522:
5521:
5519:
5516:
5512:
5505:
5504:
5500:
5498:
5497:
5493:
5492:
5489:
5486:
5483:
5478:
5475:
5472:
5471:Neoazhdarchia
5467:
5461:
5460:
5456:
5454:
5453:
5449:
5446:
5445:
5441:
5438:
5437:
5433:
5431:
5430:
5426:
5424:
5423:
5419:
5418:
5415:
5412:
5409:
5408:Azhdarchoidea
5404:
5390:
5389:
5385:
5383:
5382:
5378:
5377:
5375:
5373:
5369:
5362:
5361:
5357:
5354:
5353:
5352:Puntanipterus
5349:
5347:
5346:
5342:
5340:
5339:
5335:
5332:
5331:
5327:
5324:
5323:
5319:
5318:
5315:
5312:
5310:
5306:
5302:
5299:
5296:
5291:
5284:
5283:
5279:
5278:
5275:
5272:
5269:
5264:
5260:
5256:
5251:
5247:
5238:
5232:
5206:
5205:
5204:
5203:
5201:
5199:
5195:
5188:
5187:
5183:
5182:
5179:
5176:
5174:
5170:
5166:
5140:
5139:
5135:
5133:
5132:
5128:
5126:
5125:
5121:
5119:
5118:
5117:Beipiaopterus
5114:
5113:
5111:
5109:
5105:
5099:
5098:
5094:
5093:
5090:
5087:
5085:
5081:
5075:
5074:
5073:Tacuadactylus
5070:
5068:
5067:
5063:
5061:
5060:
5056:
5053:
5052:
5048:
5046:
5045:
5041:
5040:
5038:
5036:
5032:
5026:
5025:
5021:
5019:
5018:
5014:
5013:
5011:
5008:
5004:
4998:
4997:
4993:
4991:
4990:
4986:
4984:
4983:
4979:
4977:
4976:
4972:
4970:
4969:
4965:
4963:
4962:
4958:
4956:
4955:
4954:Forfexopterus
4951:
4949:
4948:
4947:Elanodactylus
4944:
4942:
4941:
4940:Cratonopterus
4937:
4935:
4934:
4933:Cathayopterus
4930:
4928:
4927:
4923:
4920:
4919:
4918:Ardeadactylus
4915:
4914:
4911:
4908:
4905:
4900:
4885:
4884:
4880:
4878:
4877:
4873:
4870:
4869:
4868:Ardeadactylus
4865:
4864:
4862:
4860:
4856:
4849:
4848:
4844:
4842:
4841:
4837:
4836:
4833:
4830:
4828:
4824:
4817:
4816:
4812:
4809:
4808:
4804:
4801:
4800:
4796:
4794:
4793:
4792:Cycnorhamphus
4789:
4786:
4785:
4781:
4780:
4778:
4776:
4772:
4768:
4765:
4762:
4757:
4751:
4750:
4749:Pterodactylus
4746:
4744:
4743:
4742:Diopecephalus
4739:
4738:
4735:
4732:
4729:
4724:
4713:
4712:
4708:
4705:
4704:
4700:
4698:
4697:
4693:
4690:
4689:
4685:
4684:
4682:
4680:
4676:
4670:
4669:
4668:Prejanopterus
4665:
4664:
4661:
4657:
4654:
4651:
4646:
4642:
4638:
4633:
4629:
4620:
4615:
4578:
4577:
4576:
4575:
4573:
4571:
4567:
4561:
4560:
4556:
4553:
4552:
4548:
4546:
4545:
4541:
4539:
4538:
4534:
4532:
4531:
4527:
4525:
4524:
4520:
4517:
4516:
4515:Herbstosaurus
4512:
4510:
4509:
4508:Eurolimnornis
4505:
4503:
4502:
4501:Dermodactylus
4498:
4497:
4494:
4491:
4489:
4485:
4475:
4474:
4470:
4467:
4466:
4462:
4459:
4458:
4454:
4452:
4451:
4447:
4445:
4444:
4440:
4439:
4437:
4435:
4431:
4425:
4424:
4420:
4418:
4417:
4413:
4410:
4409:
4405:
4402:
4401:
4397:
4395:
4394:
4390:
4389:
4387:
4385:
4381:
4375:
4374:
4370:
4369:
4366:
4363:
4361:
4357:
4354:
4351:
4350:Caelidracones
4346:
4340:
4339:
4335:
4332:
4331:
4327:
4326:
4323:
4320:
4317:
4312:
4294:
4293:
4292:Wukongopterus
4289:
4286:
4285:
4281:
4279:
4278:
4277:Darwinopterus
4274:
4272:
4271:
4270:Cuspicephalus
4267:
4264:
4263:
4259:
4258:
4256:
4254:
4250:
4244:
4243:
4239:
4238:
4235:
4232:
4230:
4226:
4220:
4219:
4218:Pterorhynchus
4215:
4213:
4212:
4208:
4207:
4204:
4201:
4199:
4195:
4188:
4187:
4183:
4181:
4180:
4176:
4174:
4173:
4169:
4168:
4165:
4161:
4158:
4155:
4150:
4143:
4142:
4138:
4135:
4134:
4130:
4129:
4126:
4123:
4120:
4115:
4111:
4107:
4102:
4098:
4089:
4085:
4082:
4074:
4028:
4027:
4026:
4025:
4023:
4021:
4017:
4010:
4009:
4005:
4003:
4002:
4001:Scaphognathus
3998:
3996:
3995:
3991:
3989:
3988:
3984:
3982:
3981:
3977:
3976:
3974:
3971:
3967:
3964:
3961:
3957:
3939:
3938:
3934:
3931:
3930:
3926:
3923:
3922:
3918:
3917:
3915:
3913:
3909:
3903:
3902:
3898:
3895:
3894:
3890:
3887:
3886:
3882:
3880:
3879:
3875:
3873:
3872:
3868:
3867:
3865:
3863:
3859:
3856:
3854:
3850:
3844:
3843:
3839:
3837:
3836:
3832:
3830:
3829:
3825:
3823:
3822:
3818:
3816:
3815:
3811:
3809:
3808:
3804:
3802:
3801:
3797:
3796:
3793:
3790:
3788:
3784:
3777:
3776:
3772:
3769:
3768:
3764:
3762:
3761:
3757:
3754:
3753:
3749:
3748:
3745:
3742:
3740:
3736:
3733:
3730:
3725:
3715:
3714:
3710:
3708:
3707:
3703:
3702:
3700:
3698:
3694:
3690:
3687:
3684:
3679:
3672:
3671:
3670:Eudimorphodon
3667:
3666:
3663:
3660:
3656:
3651:
3640:
3639:
3635:
3632:
3631:
3630:Peteinosaurus
3627:
3624:
3623:
3619:
3617:
3616:
3612:
3610:
3609:
3605:
3602:
3601:
3597:
3596:
3594:
3592:
3588:
3582:
3581:
3580:Herbstosaurus
3577:
3576:
3573:
3569:
3566:
3563:
3558:
3551:
3550:
3549:Peteinosaurus
3546:
3545:
3542:
3539:
3535:
3530:
3512:
3511:
3510:Eudimorphodon
3507:
3504:
3503:
3499:
3498:
3496:
3494:
3490:
3483:
3482:
3478:
3477:
3474:
3471:
3469:
3465:
3459:
3458:
3454:
3452:
3451:
3447:
3445:
3444:
3440:
3437:
3436:
3432:
3431:
3429:
3427:
3423:
3420:
3418:
3414:
3407:
3406:
3405:Preondactylia
3402:
3399:
3398:
3397:Peteinosaurus
3394:
3391:
3390:
3386:
3385:
3382:
3379:
3376:
3375:Eopterosauria
3371:
3357:
3356:
3352:
3350:
3349:
3345:
3342:
3341:
3337:
3336:
3334:
3332:
3328:
3321:
3320:
3316:
3314:
3313:
3309:
3306:
3305:
3301:
3300:
3297:
3294:
3291:
3287:
3281:
3280:
3279:Preondactylus
3276:
3274:
3273:
3269:
3268:
3266:
3264:
3263:Preondactylia
3260:
3254:
3253:
3249:
3248:
3245:
3241:
3238:
3235:
3230:
3226:
3222:
3217:
3213:
3192:
3191:
3190:
3189:
3187:
3185:
3181:
3173:
3169:
3168:
3167:
3166:
3163:
3160:
3158:
3154:
3150:
3142:
3138:
3135:
3133:
3129:
3126:
3124:
3120:
3117:
3115:
3111:
3109:
3105:
3103:
3099:
3098:
3093:
3089:
3085:
3078:
3073:
3071:
3066:
3064:
3059:
3058:
3055:
3051:
3050:
3039:
3035:
3030:
3025:
3020:
3015:
3011:
3007:
3003:
2999:
2995:
2988:
2980:
2976:
2972:
2965:
2957:
2953:
2949:
2945:
2941:
2937:
2933:
2926:
2918:
2914:
2909:
2904:
2899:
2894:
2890:
2886:
2882:
2878:
2874:
2867:
2861:
2857:
2853:
2850:
2843:
2835:
2831:
2826:
2821:
2817:
2813:
2809:
2805:
2801:
2797:
2793:
2791:
2782:
2780:
2771:
2767:
2763:
2759:
2755:
2751:
2744:
2736:
2732:
2728:
2724:
2717:
2710:
2702:
2698:
2694:
2690:
2686:
2679:
2671:
2667:
2662:
2657:
2652:
2647:
2643:
2639:
2635:
2633:
2624:
2616:
2612:
2608:
2606:
2597:
2595:
2593:
2591:
2582:
2578:
2573:
2568:
2564:
2560:
2556:
2549:
2540:
2535:
2531:
2527:
2523:
2516:
2514:
2512:
2503:
2499:
2495:
2491:
2487:
2483:
2476:
2474:
2465:
2463:
2461:
2459:
2450:
2446:
2442:
2438:
2434:
2432:
2423:
2415:
2411:
2404:
2402:
2393:
2391:
2389:
2387:
2385:
2383:
2381:
2379:
2370:
2366:
2361:
2356:
2352:
2348:
2344:
2342:
2338:
2337:Tropeognathus
2328:
2326:
2317:
2313:
2309:
2302:
2294:
2290:
2285:
2280:
2275:
2270:
2266:
2262:
2260:
2251:
2249:
2247:
2242:
2232:
2229:
2227:
2224:
2223:
2217:
2215:
2211:
2196:
2188:
2187:
2179:
2178:
2170:
2169:
2161:
2160:
2157:
2156:
2155:
2148:
2147:
2144:
2143:
2135:
2134:
2131:
2130:
2129:
2128:Maaradactylus
2122:
2121:
2118:
2117:
2114:
2113:
2112:
2111:Cearadactylus
2105:
2104:
2098:
2097:
2094:
2093:
2090:
2089:
2088:
2081:
2080:
2074:
2073:
2070:
2069:
2061:
2060:
2057:
2056:
2055:
2048:
2047:
2044:
2043:
2040:
2039:
2038:
2031:
2030:
2027:
2026:
2023:
2022:
2021:
2014:
2013:
2007:
2006:
2000:
1999:Anhanguerinae
1997:
1996:
1993:
1992:
1984:
1983:
1975:
1974:
1966:
1965:
1962:
1961:
1960:
1953:
1952:
1949:
1948:
1945:
1944:
1943:
1936:
1935:
1929:
1928:
1925:
1924:
1921:
1920:
1919:
1912:
1911:
1905:
1904:
1901:
1900:
1897:
1896:
1895:
1888:
1887:
1881:
1878:
1877:
1874:
1873:
1865:
1864:
1856:
1855:
1852:
1851:
1850:
1843:
1842:
1839:
1838:
1835:
1834:
1833:
1826:
1825:
1819:
1818:
1815:
1814:
1806:
1805:
1802:
1801:
1800:
1799:Tropeognathus
1793:
1792:
1789:
1788:
1785:
1784:
1783:
1782:Siroccopteryx
1776:
1775:
1769:
1768:
1762:
1759:
1758:
1752:
1751:Anhangueridae
1749:
1748:
1744:
1742:
1738:
1731:
1730:
1722:
1721:
1713:
1712:
1704:
1703:
1695:
1694:
1686:
1685:
1677:
1676:
1668:
1667:
1659:
1658:
1650:
1649:
1646:
1645:
1644:
1637:
1636:
1633:
1632:
1629:
1628:
1627:
1620:
1619:
1613:
1612:
1609:
1608:
1605:
1604:
1603:
1596:
1595:
1589:
1588:
1585:
1584:
1581:
1580:
1579:
1572:
1571:
1565:
1562:
1561:
1558:
1557:
1554:
1553:
1552:
1551:Tropeognathus
1545:
1544:
1538:
1535:
1534:
1531:
1530:
1522:
1521:
1518:
1517:
1516:
1509:
1508:
1505:
1504:
1501:
1500:
1499:
1492:
1491:
1485:
1484:Anhangueridae
1482:
1481:
1475:
1472:
1471:
1468:
1467:
1464:
1463:
1462:
1461:Cearadactylus
1455:
1454:
1448:
1447:
1444:
1443:
1435:
1434:
1431:
1430:
1429:
1422:
1421:
1418:
1417:
1414:
1413:
1412:
1405:
1404:
1398:
1397:
1391:
1390:
1384:
1383:
1380:
1379:
1376:
1375:
1374:
1367:
1366:
1359:
1355:
1354:
1350:
1348:
1344:
1340:
1339:
1332:
1328:
1324:
1322:
1321:Anhangueridae
1318:
1314:
1310:
1306:
1305:Siroccopteryx
1302:
1298:
1294:
1290:
1286:
1282:
1278:
1274:
1270:
1266:
1256:
1254:
1250:
1245:
1243:
1238:
1234:
1230:
1226:
1225:
1224:Tropeognathus
1220:
1213:
1212:C. fluviferox
1209:
1205:
1201:
1199:
1195:
1190:
1188:
1187:
1182:
1177:
1175:
1171:
1167:
1166:
1161:
1160:
1154:
1152:
1148:
1144:
1140:
1139:type specimen
1132:
1127:
1113:
1112:C. fluviferox
1110:
1105:
1103:
1098:
1095:
1090:
1089:
1088:C. spielbergi
1085:
1082:
1081:
1076:
1075:
1071:
1068:
1067:
1062:
1059:
1054:
1053:
1049:
1044:
1040:
1039:
1034:
1031:
1025:
1021:
1020:
1016:
1011:
1007:
1005:
1000:
997:
992:
988:
986:
981:
978:
973:
972:
967:
966:
961:
956:
954:
953:Cearadactylus
949:
945:
943:
939:
936:
935:
934:
932:
925:
920:
911:
909:
905:
901:
900:N. fluviferox
897:
896:
891:
890:C. fluviferox
887:
883:
879:
875:
874:Kem Kem Group
871:
867:
866:C. fluviferox
863:
859:
855:
851:
847:
842:
840:
839:
834:
830:
826:
822:
818:
814:
810:
809:C. sedgwickii
806:
802:
798:
793:
791:
787:
786:Siroccopteryx
783:
779:
775:
771:
767:
766:
761:
760:
755:
754:
749:
745:
741:
736:
734:
730:
726:
722:
718:
714:
712:
707:
703:
699:
695:
691:
687:
683:
682:
676:
674:
673:
668:
664:
660:
656:
652:
649:
645:
641:
640:
635:
634:Yuong-Nam Lee
630:
628:
624:
620:
616:
612:
608:
604:
600:
599:
594:
590:
586:
582:
581:O. sedgwickii
578:
574:
573:specific name
570:
566:
562:
561:Wealden Group
558:
554:
550:
549:
545:of the genus
544:
540:
535:
533:
529:
525:
517:
513:
509:
500:
498:
494:
493:
492:Tropeognathus
488:
484:
480:
476:
472:
468:
464:
460:
456:
455:Anhangueridae
452:
449:
445:
441:
440:
421:
415:
412:
411:
410:
409:
406:
405:C. fluviferox
390:
388:
384:
378:
376:
372:
371:
370:
369:
366:
352:
351:
344:
341:
340:
339:
338:
331:
330:
322:
319:
315:
308:
301:
300:
299:C. fluviferox
293:
289:
282:
281:
274:
273:
271:
266:
259:
258:
250:
247:
243:
238:
233:
232:
225:
222:
221:
218:
212:
209:
206:
205:
202:
196:
193:
192:
189:
183:
180:
179:
176:
173:
170:
169:
166:
163:
160:
159:
156:
153:
150:
149:
144:
139:
135:
131:
127:
123:
119:
114:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
39:
36:140–136
32:
26:
22:
19:
7802:Paleontology
7702:
7666:
7612:
7607:Ludodactylus
7605:
7598:
7591:
7583:
7575:
7568:
7548:
7540:
7535:Nicorhynchus
7533:
7527:
7526:
7519:
7498:
7491:
7483:
7475:
7467:
7459:
7451:
7421:
7414:
7406:Hamipteridae
7392:
7384:
7350:
7342:
7334:
7314:
7306:
7301:Aetodactylus
7299:
7264:
7256:
7249:
7241:
7235:Ludodactylus
7233:
7225:
7217:
7209:
7202:
7201:
7193:
7187:Aetodactylus
7185:
7164:
7156:
7148:
7140:
7132:
7125:
7118:
7075:
7068:
7044:
7037:
7030:
7023:
6997:Liaoxipterus
6995:
6988:
6968:
6961:
6954:
6947:
6925:Mimodactylus
6923:
6916:
6909:
6888:
6881:
6874:
6867:
6843:
6836:
6811:
6803:Lanceodontia
6784:
6776:
6771:Lonchodectes
6769:
6761:
6753:
6733:
6726:
6719:
6672:
6665:
6658:
6651:
6631:
6624:
6617:
6592:
6585:
6560:
6552:
6545:
6539:Ornithostoma
6537:
6530:
6522:
6493:
6433:
6347:
6339:
6332:
6325:
6318:
6311:
6306:Eurazhdarcho
6304:
6297:
6290:
6283:
6263:
6256:
6249:
6241:
6234:Azhdarchinae
6220:
6212:
6205:
6197:
6190:
6182:
6175:Azhdarchidae
6161:
6154:
6146:
6118:
6111:
6104:
6083:
6075:
6070:Meilifeilong
6068:
6061:
6053:
6045:
6037:
6013:
5988:
5980:
5972:
5964:
5956:
5949:
5927:
5920:
5913:
5905:
5855:
5848:
5828:
5821:
5814:
5806:
5782:
5774:
5766:
5758:
5750:
5726:
5719:
5712:
5707:Huaxiapterus
5705:
5698:
5692:Eopteranodon
5690:
5682:
5676:Afrotapejara
5674:
5667:Sinopterinae
5656:Afrotapejara
5654:
5630:
5623:
5592:
5585:
5578:
5570:
5562:
5555:
5547:
5539:
5531:
5523:
5501:
5496:Bennettazhia
5494:
5459:Ornithostoma
5457:
5450:
5442:
5434:
5427:
5420:
5386:
5379:
5358:
5350:
5345:Ordosipterus
5343:
5336:
5328:
5320:
5295:Tapejaroidea
5280:
5184:
5138:Pterodaustro
5136:
5129:
5122:
5115:
5095:
5071:
5064:
5057:
5051:Huanhepterus
5049:
5044:Gnathosaurus
5042:
5024:Moganopterus
5022:
5015:
4996:Pterofiltrus
4994:
4989:Petrodactyle
4987:
4980:
4975:Liaodactylus
4973:
4968:Kepodactylus
4966:
4959:
4952:
4945:
4938:
4931:
4924:
4916:
4883:Huanhepterus
4881:
4874:
4866:
4845:
4840:Aerodactylus
4838:
4813:
4807:Petrodactyle
4805:
4797:
4790:
4782:
4747:
4740:
4709:
4701:
4694:
4686:
4666:
4557:
4549:
4542:
4535:
4528:
4523:Kryptodrakon
4521:
4513:
4506:
4499:
4471:
4465:Jeholopterus
4463:
4455:
4448:
4441:
4421:
4414:
4408:Jeholopterus
4406:
4398:
4393:Anurognathus
4391:
4373:Mesadactylus
4371:
4336:
4328:
4290:
4282:
4275:
4268:
4260:
4240:
4216:
4209:
4198:Darwinoptera
4184:
4177:
4170:
4139:
4131:
4077:
4006:
3999:
3992:
3985:
3978:
3935:
3929:Nesodactylus
3927:
3921:Cacibupteryx
3919:
3901:Sericipterus
3899:
3891:
3883:
3876:
3869:
3840:
3833:
3828:Nesodactylus
3826:
3819:
3812:
3807:Cacibupteryx
3805:
3800:Bellubrunnus
3798:
3773:
3765:
3758:
3750:
3711:
3704:
3668:
3655:Lonchognatha
3636:
3628:
3620:
3613:
3606:
3598:
3578:
3547:
3508:
3500:
3479:
3455:
3448:
3443:Pachagnathus
3441:
3433:
3403:
3395:
3389:Austriadraco
3387:
3353:
3348:Austriadraco
3346:
3338:
3317:
3310:
3302:
3277:
3270:
3250:
3136:
3127:
3118:
3047:
3001:
2997:
2987:
2970:
2964:
2939:
2935:
2931:
2925:
2883:(1): 17875.
2880:
2876:
2866:
2851:
2848:
2842:
2802:(1): 13454.
2799:
2795:
2789:
2753:
2749:
2743:
2726:
2722:
2709:
2692:
2688:
2684:
2678:
2641:
2637:
2631:
2623:
2614:
2610:
2604:
2562:
2558:
2548:
2529:
2525:
2485:
2481:
2472:
2440:
2436:
2430:
2422:
2413:
2409:
2400:
2350:
2346:
2340:
2336:
2316:the original
2311:
2301:
2264:
2258:
2209:
2207:
2204:Paleobiology
2194:
2152:
2151:
2126:
2125:
2109:
2108:
2085:
2084:
2054:Ludodactylus
2052:
2051:
2035:
2034:
2018:
2017:
1957:
1956:
1942:Nicorhynchus
1940:
1939:
1917:
1916:
1915:
1892:
1891:
1847:
1846:
1830:
1829:
1797:
1796:
1780:
1779:
1740:
1739:
1641:
1640:
1624:
1623:
1600:
1599:
1577:
1576:
1575:
1549:
1548:
1513:
1512:
1496:
1495:
1459:
1458:
1428:Ludodactylus
1426:
1425:
1409:
1408:
1371:
1370:
1346:
1342:
1341:
1330:
1316:
1313:C. wadleighi
1312:
1311:to refer to
1308:
1304:
1300:
1296:
1292:
1288:
1284:
1276:
1272:
1269:C. wadleighi
1268:
1264:
1262:
1255:of England.
1248:
1246:
1241:
1236:
1222:
1218:
1216:
1211:
1197:
1191:
1184:
1180:
1178:
1173:
1169:
1163:
1157:
1155:
1150:
1146:
1142:
1136:
1130:
1111:
1100:
1096:
1086:
1078:
1072:
1064:
1060:
1052:C. wadleighi
1050:
1042:
1036:
1030:nomen dubium
1029:
1023:
1017:
1009:
1002:
999:C. sedgwicki
998:
990:
983:
979:
969:
963:
951:
947:
942:type species
937:
930:
928:
923:
907:
904:Nicorhynchus
903:
899:
895:Nicorhynchus
893:
889:
885:
865:
861:
857:
854:nomina dubia
853:
849:
845:
843:
838:Lonchodectes
836:
832:
828:
824:
821:C. sedgwicki
820:
816:
812:
808:
804:
800:
794:
789:
785:
781:
777:
774:C. wadleighi
773:
769:
763:
757:
751:
747:
743:
739:
737:
732:
728:
724:
720:
716:
709:
701:
697:
679:
677:
670:
666:
663:C. wadleighi
662:
658:
654:
637:
631:
627:Criorhynchus
626:
622:
618:
614:
610:
606:
602:
596:
589:type species
584:
580:
576:
552:
546:
539:Richard Owen
536:
531:
527:
521:
515:
490:
486:
462:
438:
437:
436:
419:
413:
404:
403:Synonyms of
387:Nicorhynchus
385:
381:Seeley, 1870
373:
364:
363:Synonyms of
348:
342:
306:
297:
278:
256:
255:
246:Type species
230:
229:
207:
129:
24:
18:
7727:Wikispecies
7493:Thapunngaka
7453:Amblydectes
7372:Anhangueria
7336:Aussiedraco
7070:Boreopterus
6845:Lonchodraco
6778:Lonchodraco
6721:Aussiedraco
6667:Nyctosaurus
6524:Bogolubovia
6192:Bogolubovia
6063:Lacusovagus
6055:Eoazhdarcho
6015:Eoazhdarcho
5974:Leptostomia
5966:Keresdrakon
5841:Caiuajarina
5808:Bakonydraco
5776:Lacusovagus
5768:Keresdrakon
5739:Tapejarinae
5700:Huaxiadraco
5684:Bakonydraco
5643:Tapejaridae
5572:Leptostomia
5564:Lacusovagus
5557:Kariridraco
5503:Keresdrakon
5444:Leptostomia
5436:Keresdrakon
5097:Ctenochasma
5059:Lusognathus
4982:Otogopterus
4637:Lophocratia
4570:Lophocratia
4537:Pangupterus
4473:Sinomacrops
3814:Dorygnathus
3683:Novialoidea
3638:Rhamphinion
3615:Dimorphodon
3290:Caviramidae
3234:Pterosauria
3221:Pterosauria
3184:Pterosauria
3123:Archosauria
3084:Pterosauria
3004:(1): 5293.
1741:Topology 2:
1358:Anhangueria
1343:Topology 1:
1121:Description
1074:C. piscator
1038:C. robustus
1012:Owen, 1859
965:C. piscator
948:C. ligabuei
717:C. robustus
706:David Unwin
704:. In 2002,
565:East Sussex
475:Valanginian
350:nomen nudum
188:Pterosauria
7824:Categories
7570:Anhanguera
7461:Ferrodraco
7416:Hamipterus
7344:Barbosania
7219:Ferrodraco
7211:Draigwenia
7046:Hamipterus
7025:Barbosania
6970:Nurhachius
6838:Ikrandraco
6813:Draigwenia
6763:Ikrandraco
6728:Serradraco
6674:Volgadraco
6562:Volgadraco
6554:Tethydraco
6547:Pteranodon
6405:see below↓
6299:Cryodrakon
6222:Volgadraco
6214:Tethydraco
6113:Jidapterus
6077:Microtuban
5915:Microtuban
5816:Europejara
5795:Tapejarini
5784:Vectidraco
5721:Sinopterus
5338:Noripterus
5207:see below↓
5131:Gegepterus
5124:Eosipterus
4579:see below↓
4559:Wenupteryx
4450:Cascocauda
4029:see below↓
3457:Yelaphomte
3193:see below↓
3114:Sauropsida
2632:Anhanguera
2416:: 219–228.
2410:Zitteliana
2312:BBC Nature
2237:References
2154:Anhanguera
1849:Ferrodraco
1626:Ferrodraco
1515:Anhanguera
1170:Anhanguera
1159:Anhanguera
1055:Lee, 1994
980:C. cuvieri
733:Anhanguera
711:Anhanguera
522:Like many
512:Lithograph
483:Cenomanian
262:Owen, 1874
194:Suborder:
7521:Aerodraco
6963:Luchibang
6911:Haopterus
6633:Simurghia
6626:Epapatelo
6594:Cretornis
6532:Dawndraco
6258:Azhdarcho
6251:Albadraco
6243:Aerotitan
6163:Alanqidae
5941:Alanqidae
5907:Cretornis
5857:Torukjara
5850:Caiuajara
5587:Tupuxuara
5525:Aerotitan
5429:Inabtanin
5017:Feilongus
4416:Luopterus
4133:Allkaruen
3760:Klobiodon
3600:Allkaruen
3435:Caviramus
3312:Caviramus
3100:Kingdom:
2956:209595986
2644:: e3285.
2581:1867-6812
2502:134439172
2488:: 77–88.
2214:piscivore
1894:Aerodraco
1345:Pentland
1237:C. capito
1097:C. capito
1006:sedgwicki
924:C. capito
886:C. capito
868:from the
813:C. capito
805:C. capito
684:from the
555:based on
451:pterosaur
365:C. capito
280:C. capito
161:Kingdom:
155:Eukaryota
7712:Wikidata
7593:Guidraco
7477:Mythunga
7469:Haliskia
7243:Mythunga
7227:Guidraco
7158:Haliskia
6735:Unwindia
6085:Xericeps
5990:Xericeps
5922:Xericeps
5823:Tapejara
5594:Xericeps
5549:Banguela
5322:Banguela
4551:Samrukia
4211:Ceoptera
3108:Chordata
3106:Phylum:
3102:Animalia
3038:33116130
2917:31784545
2834:31582757
2770:84617119
2670:28484676
2369:23538956
2293:23794925
2220:See also
2037:Guidraco
1832:Mythunga
1643:Mythunga
1373:Guidraco
1349:(2019).
955:ligabuei
713:piscator
557:holotype
537:In 1874
518:holotype
497:wingspan
418:(Jacobs
318:Synonyms
175:Chordata
171:Phylum:
165:Animalia
151:Domain:
126:Holotype
7788:Portals
7760:1165136
7747:4818490
7718:Q286251
6619:Alcione
5728:Wightia
3112:Class:
3029:7595196
3006:Bibcode
2908:6884559
2885:Bibcode
2854:: 4940
2825:6776501
2804:Bibcode
2695:: 1–9.
2661:5420195
2284:3689139
2265:ZooKeys
1360:
1233:England
987:cuvieri
878:Morocco
872:of the
688:of the
569:England
514:of the
471:England
422:, 2019)
305:Jacobs
290:, 1870)
223:Genus:
181:Order:
105:↓
6184:Alanqa
5951:Alanqa
5533:Alanqa
4141:Sordes
4008:Sordes
3036:
3026:
2954:
2915:
2905:
2832:
2822:
2768:
2668:
2658:
2579:
2500:
2367:
2291:
2281:
2195:
1356:
1347:et al.
1297:et al.
1285:et al.
1104:capito
858:et al.
694:Brazil
644:Albian
636:named
479:Albian
420:et al.
389:capito
377:capito
309:, 2019
307:et al.
288:Seeley
239:, 1874
7773:55642
7755:IRMNG
5282:Piksi
3878:Dearc
3137:Clade
3128:Clade
3119:Clade
2952:S2CID
2766:S2CID
2719:(PDF)
2638:PeerJ
2498:S2CID
2478:(PDF)
2406:(PDF)
906:from
715:with
651:Texas
444:genus
442:is a
208:Clade
7742:GBIF
3170:see
3034:PMID
2913:PMID
2830:PMID
2666:PMID
2577:ISSN
2559:PalZ
2365:PMID
2339:cf.
2289:PMID
1303:and
1267:and
1162:and
1137:The
888:and
807:and
727:and
646:age
579:and
481:and
237:Owen
44:PreꞒ
3024:PMC
3014:doi
2975:doi
2944:doi
2903:PMC
2893:doi
2856:doi
2820:PMC
2812:doi
2758:doi
2754:103
2731:doi
2727:126
2697:doi
2656:PMC
2646:doi
2615:125
2567:doi
2534:doi
2490:doi
2445:doi
2355:doi
2279:PMC
2269:doi
1231:of
1141:of
876:of
841:).
799:to
768:to
692:in
563:of
469:of
446:of
7826::
7770::
7757::
7744::
7729::
7714::
3139::
3130::
3121::
3032:.
3022:.
3012:.
3002:11
3000:.
2996:.
2950:.
2940:33
2938:.
2911:.
2901:.
2891:.
2879:.
2875:.
2828:.
2818:.
2810:.
2798:.
2794:.
2778:^
2764:.
2752:.
2725:.
2721:.
2693:34
2691:.
2664:.
2654:.
2640:.
2636:.
2613:.
2609:.
2589:^
2575:.
2563:97
2561:.
2557:.
2532:.
2530:65
2528:.
2524:.
2510:^
2496:.
2486:95
2484:.
2480:.
2457:^
2441:13
2439:.
2435:.
2414:28
2412:.
2408:.
2377:^
2363:.
2351:85
2349:.
2345:.
2324:^
2310:.
2287:.
2277:.
2263:.
2245:^
864:,
835:(=
831:"
803::
735:.
675:.
629:.
567:,
499:.
302:?
283:?
210::
94:Pg
38:Ma
33:,
7790::
7588:?
7580:?
7545:?
7513:?
7488:?
7480:?
7472:?
7464:?
7456:?
7445:?
7408:?
7397:?
7389:?
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7339:?
7311:?
7284:?
7269:?
7261:?
7246:?
7238:?
7230:?
7222:?
7214:?
7206:?
7198:?
7190:?
7169:?
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6893:?
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6789:?
6781:?
6766:?
6758:?
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6597:?
6565:?
6557:?
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6225:?
6217:?
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6088:?
6080:?
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5932:?
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5811:?
5779:?
5771:?
5763:?
5755:?
5695:?
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5679:?
5617:?
5597:?
5575:?
5567:?
5552:?
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5536:?
5528:?
5517:?
5506:?
5447:?
5439:?
5363:?
5355:?
5333:?
5325:?
5285:?
5189:?
5054:?
5009:?
4921:?
4886:?
4871:?
4850:?
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4802:?
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4714:?
4706:?
4691:?
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4468:?
4460:?
4411:?
4403:?
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4287:?
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4011:?
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3755:?
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3641:?
3633:?
3625:?
3603:?
3552:?
3536:?
3505:?
3484:?
3438:?
3408:?
3400:?
3392:?
3343:?
3322:?
3307:?
3292:?
3076:e
3069:t
3062:v
3040:.
3016::
3008::
2981:.
2977::
2958:.
2946::
2919:.
2895::
2887::
2881:9
2858::
2852:9
2836:.
2814::
2806::
2800:9
2788:"
2772:.
2760::
2737:.
2733::
2703:.
2699::
2672:.
2648::
2642:5
2630:"
2583:.
2569::
2542:.
2536::
2504:.
2492::
2451:.
2447::
2403:"
2371:.
2357::
2295:.
2271::
1035:?
962:?
946:?
473:(
353:)
295:†
286:(
276:†
253:†
227:†
214:†
198:†
185:†
99:N
89:K
84:J
79:T
74:P
69:C
64:D
59:S
54:O
49:Ꞓ
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