870:
450:
461:
140:
3144:
624:
3447:
1286:
113:
1885:, which indicates that the group had spread significantly during their 70 million year existence. Pterygotioids appear to have been relatively abundant throughout the Silurian and Devonian world, with unusually (in terms of eurypterids, most groups being absent entirely) large numbers recovered from the ancient continent of
611:. These appendages are the only ones that appear before the mouth and take the form of small pincers used to feed in most eurypterid groups. This function is retained in the more basal Hughmilleriidae and Slimonidae, but pterygotid chelicerae were large and long with strong and well developed teeth on specialized
1315:
Though the
Pterygotidae are accepted to clearly represent the most derived group within the pterygotioid superfamily, there has been an ongoing debate on whether the hughmilleriids or the slimonids are the most closely related to the pterygotids, and thus also which of the two families is the most
919:
Pterygotioidea is the most derived superfamily of the suborder and contains over 50 species (the exact number obscured by dubious species and possible synonyms), which accounts for more than a fifth of the approximately 250 known eurypterid species. The closest sister group of the
Pterygotioidea,
403:
Among all currently recognized eurypterid clades, the
Pterygotioidea is the most diverse, containing over 50 species in 10 genera. With the number of recognized eurypterid species being around 250, pterygotioids account for more than a fifth of all known eurypterid species. Though the group only
1935:
It remains unclear whether pterygotioids were capable of crossing oceans on a regular basis or if species recovered outside of
Laurentia, Baltica and Avalonia represent isolated occurrences. Pterygotioids were, like the related adelophthalmoids, excellent swimmers which might help explain the
431:
Though several characteristic and diagnostic traits can be established for each of the families included within the
Pterygotioidea, the group as a whole is primarily joined by the shared features of marginal eyes, that their compound eyes are placed near or on the margin of the
1351:
The cladogram presented below, derived from a 2007 study by researcher O. Erik Tetlie, showcases the interrelationships between the pterygotioid eurypterids. Whilst
Slimonidae and Pterygotidae form monophyletic (and thus valid) groups, Hughmilleriidae has been recovered as
399:
being the smallest of all pterygotioids at just 6 centimetres (2.4 in) in length, later members of the group, particularly in the
Pterygotidae, would become the largest known arthropods to ever exist with several genera surpassing 2 metres (6.6 ft) in length.
672:, meaning "winged one". Agassiz mistakenly believed that the fossils were the remains of a large fish. Agassiz first recognized the true nature of the fossils as arthropod remains five years later in 1844 after having examined more complete fossils recovered in the
1928:. With the group already being widespread at this seemingly early stage in their evolution, it is difficult to pinpoint their exact geographical point of origin. The closest relatives and sister clade of the pterygotioids, the superfamily
2338:
The
Silurian System, Founded on Geological Researches in the Counties of Salop, Hereford, Radnor, Montgomery, Caermarthen, Brecon, Pembroke, Monmouth, Gloucester, Worcester, and Stafford: With Descriptions of the Coalfields and Overlying
2029:
Olive, Sébastien; Pradel, Alan; Martinez-Pérez, Carlos; Janvier, Philippe; Lamsdell, James C.; Gueriau, Pierre; Rabet, Nicolas; Duranleau-Gagnon, Philippe; Cardenas-Rozo, Andres L.; Zapata
Ramirez, Paula A.; Botella, Héctor (2019).
2526:
1337:, combined with the fact that the genus shares certain characteristics with basal adelophthalmids (in particular the triangular anterior margin of the carapace), places it as the most basal genus in the superfamily.
594:(the posteriormost segment of the body) was expanded and flattened, often with a spike protruding from its end. The telsons of the Hughmilleriidae were not flattened, instead being lanceolate (in the shape of a
1341:
also possessed far more gnathobasic (of the gnathobase, an appendage used in feeding) teeth than any other pterygotioid, possessing 18-20 whilst more derived pterygotioids (including related hughmilleriid
923:
The cladogram below is simplified from 2007 study by O. Erik Tetlie, showcasing the position of the pterygotioids within the suborder
Eurypterina. Placement of Diploperculata follows Lamsdell
1932:, originated in Laurentia. Modern researchers assume that the case would be the same with the pterygotioids, which are thought to have originated within, or in close proximity to, Laurentia.
404:
existed for around 70 million years and during a time when most continents were separated by large expanses of water (in contrast with previous and later periods of time when there had been
579:(abdomen). The appendages were attached to the prosoma, and were characterized in pterygotids and slimonids by being small and slender and lacking spines. In contrast, the more
412:
of all eurypterid groups. Their fossils have been recovered from Europe, Africa, North America, South America, Asia and Australia, with the earliest remains being from the
2322:
2668:
2676:
861:
into a pterygotioid family of its own as it was considered distinct enough from other hughmilleriids to warrant a separate family, the Slimonidae.
816:
would later be raised to its own genus. American paleontologist Erik N. Kjellesvig-Waering emended the Pterygotidae in 1951, referring the genera
1360:
and does thus not form an actually valid scientific grouping. Nevertheless, the family is retained and routinely used by eurypterid researchers.
1845:
Pterygotioid fossils have also been recovered from other parts of the world where fossils of other eurypterid groups are absent, including
741:. Its describer, the American geologist Clifton J. Sarle, considered the genus to represent an intermediate form between the more basal
2615:"A new Ordovician eurypterid (Arthropoda: Chelicerata) from southeast Turkey: Evidence for a cryptic Ordovician record of Eurypterida"
857:
and creating the superfamily Pterygotioidea, containing Hughmilleriidae and Pterygotidae. The same year, Novojilov also reclassified
1826:. During the Silurian and Devonian periods, when pterygotioids were alive, these regions and continents were part of the continents
900:
suborder of eurypterids. The infraorder Diploperculata contains the four most derived superfamilies of eurypterine eurypterids;
3223:
3213:
2735:
2209:
920:
Adelophthalmoidea, also contains a large amount of species, over 40, and is the second most diverse eurypterid superfamily.
3545:
1326:(especially within the appendages) and of more derived pterygotid eurypterids, revealing that Slimonidae was the closest
3175:
2036:
2032:"New insights into Late Devonian vertebrates and associated fauna from the Cuche Formation (Floresta Massif, Colombia)"
2425:
Kjellesvig-Waering, Erik N. (1964). "A Synopsis of the Family Pterygotidae Clarke and Ruedemann, 1912 (Eurypterida)".
1978:
Tetlie, O. Erik; Briggs, Derek E. G. (2009-09-01). "The origin of pterygotid eurypterids (Chelicerata: Eurypterida)".
3190:
2542:
3555:
853:
raised the Eurypteracea and Pterygotidae to subordinal and superfamily status, Eurypteracea becoming the suborder
3434:
3218:
3160:
1950:
2701:
3550:
3208:
3198:
3152:
2225:
Tollerton, V. P. (1989). "Morphology, taxonomy, and classification of the order Eurypterida Burmeister, 1843".
2091:"Cope's rule and Romer's theory: patterns of diversity and gigantism in eurypterids and Palaeozoic vertebrates"
3170:
3165:
3203:
3180:
139:
568:
850:
2836:
2728:
2279:
Plotnick, Roy E.; Baumiller, Tomasz K. (1988-01-01). "The pterygotid telson as a biological rudder".
2138:
1945:
505:
eurypterids by the placement of their eyes, being located near or in some cases on the margin of the
409:
3540:
3503:
1925:
1561:
587:
521:
127:
3517:
3465:
3143:
846:
711:
533:
336:
2575:. In Orlov, J. A. (ed.): Osnovy Paleontologii - volume 7, 404-423. Akademii Nauk SSSR, Moscow.
1936:
intercontinental dispersal patterns and wide-ranging distribution seen in both superfamilies.
724:
most apparently by the lack of large cheliceral claws, otherwise a defining characteristic of
3512:
1357:
580:
424:. The exact geographical origin of the group remains unknown, but is thought to have been in
371:
913:
509:(the "head" plate). Pterygotioids ranged in size from small eurypterids, the smallest being
3450:
2721:
2685:
2626:
2589:
2378:
2288:
2234:
2045:
1987:
493:
Pterygotioid eurypterids, whose fossils are recovered in deposits ranging in age from the
8:
2369:
Tetlie, O. Erik (2006). "Eurypterida (Chelicerata) from the Welsh Borderlands, England".
1839:
1537:
688:
628:
247:
2689:
2630:
2382:
2292:
2238:
2049:
1991:
2554:
2527:"Pterygotids (Chelicerata; Eurypterida) from the Silurian Vernon Formation of New York"
2481:
2434:
2402:
2300:
2258:
2180:
2153:
2125:
2071:
2011:
1479:
869:
291:
134:
122:
916:(the structure that contains the genital appendage) is made up of two fused segments.
3039:
2972:
2964:
2614:
2558:
2546:
2485:
2473:
2394:
2304:
2250:
2185:
2130:
2112:
2075:
2031:
2003:
1999:
1929:
1079:
1057:
1035:
905:
901:
761:
673:
367:
225:
2406:
2262:
2090:
2015:
1838:(most of eastern continental North America). It is around these continents, and the
449:
3379:
3093:
3085:
3075:
3067:
3047:
3018:
2934:
2883:
2693:
2634:
2538:
2465:
2386:
2336:
2296:
2242:
2175:
2167:
2120:
2104:
2061:
2053:
1995:
1917:
988:
765:
754:
738:
302:
229:
2057:
3408:
3254:
3111:
3052:
3008:
2911:
2898:
2697:
2158:
2095:
1455:
1344:
1129:
909:
561:
375:
270:
2500:
842:
311:
3488:
3324:
3282:
3240:
3030:
2990:
2982:
2906:
1815:
1022:
893:
494:
460:
413:
405:
359:
207:
28:
2390:
2246:
3534:
3331:
3303:
3296:
3268:
3247:
3000:
2939:
2929:
2921:
2875:
2865:
2857:
2813:
2777:
2638:
2550:
2477:
2398:
2308:
2254:
2116:
2007:
1898:
1874:
1823:
1819:
1010:
737:
was created based on fossils discovered in the Pittsford Shale Member of the
657:
612:
498:
352:
253:
70:
32:
2454:"III. On the Occurrence of Fossils in the Old Red Sandstone of Westmoreland"
845:
considered the pterygotid clade to represent a family within the eurypterid
3289:
3121:
3057:
3013:
2888:
2469:
2189:
2171:
2134:
2108:
1862:
1718:
1701:
1677:
1653:
1506:
1492:
1422:
1405:
1381:
1353:
1327:
1318:
1307:
1176:
1105:
875:
830:
812:
733:
704:
623:
549:
511:
479:
395:
383:
363:
282:
238:
3497:
3386:
3363:
3261:
2955:
2848:
2786:
2768:
1811:
1290:
975:
954:
934:
897:
854:
640:
576:
553:
529:
502:
421:
181:
45:
2501:"A new eurypterid fauna from the base of the Salina of western New York"
834:, to their own family within the Eurypteracea, Hughmilleriidae, leaving
3352:
3338:
3317:
3310:
3275:
3116:
2804:
2795:
2744:
2438:
2211:
Treatise on Invertebrate Paleontology, Part P Arthropoda 2, Chelicerata
2066:
1906:
1602:
1585:
1295:
692:
649:
633:
608:
600:
557:
483:
470:
387:
379:
356:
340:
331:
327:
276:
194:
112:
90:
55:
1842:(western and central Europe), that pterygotioids are the most common.
532:, and other arthropods in general, pterygotioid eurypterids possessed
3429:
3413:
3403:
2762:
1902:
1846:
1835:
1285:
791:
778:
665:
661:
598:
or spike) and similar to those of more primitive eurypterids such as
516:
425:
344:
171:
151:
95:
39:
3459:
2453:
3482:
3372:
3345:
1921:
1886:
1831:
1301:
1153:
716:
696:
677:
556:
was covered in a scale-like ornamentation but it was smooth within
506:
474:
433:
417:
85:
80:
65:
60:
50:
2669:"Distribution and dispersal history of Eurypterida (Chelicerata)"
1882:
1878:
1858:
1850:
1827:
648:
The first pterygotioid fossils to be uncovered were those of the
572:
541:
537:
100:
75:
2028:
1810:
Eurypterids are most commonly recovered from fossil deposits in
2756:
2543:
10.1666/pleo0022-3360(2007)081[0725:PEFTSV]2.0.CO;2
1890:
591:
545:
161:
2713:
1920:
of the Silurian and appear around the same time in Laurentia,
525:
which might have reached lengths of 2.6 metres (8.5 ft).
515:
at just 6 centimetres (2.4 in) in length, to the largest
2348:
as "winged fish", but there is no "fish" element in the name.
2152:
Braddy, Simon J.; Poschmann, Markus; Tetlie, O. Erik (2007).
1894:
1870:
1866:
1854:
595:
2613:
Lamsdell, James; Hoşgör, Izzet; Selden, Paul (2013-01-31).
1322:, a primitive pterygotid that clearly combines features of
714:
erected a new genus to contain the species. The new genus,
699:, were soon realized to be distinct from other species of
1912:
The earliest known pterygotioids, belonging to the genus
1316:
basal. This debate was resolved with the description of
1834:(Germany, Britain, parts of eastern North America) and
691:. The fossils referred to this species, recovered from
590:
members of the group, Slimonidae and Pterygotidae, the
583:
Hughmilleriids did possess spines on their appendages.
2590:"A summary list of fossil spiders and their relatives"
720:, could be differentiated from other known species of
2151:
393:
Though the more primitive hughmilleriids were small,
2612:
883:
is the most basal known genus in the Pterygotioidea.
668:, described the fossils in 1839 and named the genus
892:Pterygotioids are classified within the infraorder
536:bodies and jointed appendages (limbs) covered in a
2424:
2677:Palaeogeography, Palaeoclimatology, Palaeoecology
2587:
2278:
607:Like other chelicerates, pterygotioids possessed
3532:
2458:Transactions of the Edinburgh Geological Society
849:"Eurypteracea". In 1962, Russian paleontologist
841:In 1955, Norwegian paleontologist and geologist
2154:"Giant claw reveals the largest ever arthropod"
760:The family Pterygotidae was erected in 1912 by
2088:
2729:
2588:Dunlop, J. A.; Penney, D.; Jekel, D. (2018).
2089:Lamsdell, James C.; Braddy, Simon J. (2009).
386:which were equipped with robust and powerful
2524:
1977:
638:" in 1852. This species is now considered a
2525:Ciurca, Samuel J.; Tetlie, O. Erik (2007).
2344:Note that this work incorrectly translates
1333:The lack of ornamentation in the telson of
790:had also been designated as containing two
2736:
2722:
111:
2451:
2334:
2224:
2179:
2124:
2065:
567:The chelicerate body is divided into two
1284:
912:, united by the shared feature that the
868:
622:
2662:
2660:
2658:
2656:
2654:
2652:
2650:
2648:
2583:
2581:
2452:Nicholson, Henry Alleyne (1868-01-01).
2207:
1973:
1971:
1969:
1967:
1965:
3533:
2666:
2420:
2418:
2416:
2368:
2364:
2362:
2360:
2358:
2356:
2354:
1293:of each of the pterygotioid families,
618:
501:, can be distinguished from all other
3464:
3463:
2717:
2608:
2606:
2520:
2518:
2498:
2274:
2272:
2208:Størmer, Leif (1955). "Merostomata".
2145:
1280:
887:
768:to constitute a group for the genera
2645:
2578:
2321:Clarke, J. K., Ruedemann R. (1912) "
2203:
2201:
2199:
1962:
370:eurypterids. The group includes the
2413:
2351:
2022:
408:), the Pterygotioidea had the most
13:
2603:
2515:
2492:
2335:Murchison, Roderick Impey (1839).
2328:
2301:10.1111/j.1502-3931.1988.tb01746.x
2269:
2218:
2037:Journal of Vertebrate Paleontology
1830:(Scandinavia and Eastern Europe),
16:Extinct superfamily of eurypterids
14:
3567:
2315:
2196:
2082:
864:
3446:
3445:
3142:
2342:. Albemarle Street. p. 606.
2000:10.1111/j.1475-4983.2009.00907.x
519:to ever live, the largest being
459:
448:
138:
43:
3435:Timeline of eurypterid research
2743:
2565:
2445:
1951:Timeline of eurypterid research
1805:
710:) and that same year geologist
128:Senckenberg Museum of Frankfurt
2599:. Natural History Museum Bern.
2505:New York State Museum Bulletin
838:as the sole pterygotid genus.
439:
343:, an extinct group of aquatic
1:
2058:10.1080/02724634.2019.1620247
1956:
378:, the larger and specialized
335:, meaning "winged one") is a
2698:10.1016/j.palaeo.2007.05.011
326:(the name deriving from the
7:
3546:Paleozoic first appearances
2323:The Eurypterida of New York
1939:
552:, the outer surface of the
10:
3572:
2499:Sarle, Clifton J. (1902).
1719:Erettopterus waylandsmithi
1678:Erettopterus serricaudatus
851:Nestor Ivanovich Novojilov
3472:
3443:
3422:
3396:
3362:
3232:
3189:
3151:
3140:
3102:
3084:
3066:
3038:
3027:
2999:
2981:
2963:
2952:
2920:
2897:
2874:
2856:
2845:
2837:List of eurypterid genera
2829:
2822:
2751:
2391:10.1017/S0016756806002536
2247:10.1017/S0022336000041275
1946:List of eurypterid genera
1715:
1698:
1691:
1674:
1667:
1650:
1643:
1599:
1586:Acutiramus macrophthalmus
1582:
1575:
1558:
1551:
1534:
1527:
1520:
1503:
1496:
1476:
1469:
1452:
1445:
1419:
1402:
1395:
1378:
1371:
1174:
1167:
1150:
1143:
1126:
1119:
1102:
1095:
1077:
1070:
1055:
1048:
1033:
1026:
1008:
1001:
986:
979:
952:
945:
938:
908:, Pterygotioidea and the
410:cosmopolitan distribution
297:
290:
266:
261:
244:
237:
135:Scientific classification
133:
119:
110:
23:
2667:Tetlie, O. Erik (2007).
2639:10.1016/j.gr.2012.04.006
1507:Ciurcopterus ventricosus
1348:) possessed only 12–13.
571:(sections); the frontal
3556:Arthropod superfamilies
2531:Journal of Paleontology
2427:Journal of Paleontology
2227:Journal of Paleontology
1840:Rheno-Hercynian Terrane
1702:Erettopterus osiliensis
1562:Jaekelopterus rhenaniae
1456:Herefordopterus banksii
1423:Hughmilleria shawangunk
522:Jaekelopterus rhenaniae
2470:10.1144/transed.1.1.15
2172:10.1098/rsbl.2007.0491
2109:10.1098/rsbl.2009.0700
1312:
1311:, compared to a human.
884:
828:, and the newly named
645:
3551:Paleozoic extinctions
3513:Paleobiology Database
2139:Supplemental material
1406:Hughmilleria socialis
1358:phylogenetic analyses
1330:of the Pterygotidae.
1289:Largest sizes of the
1288:
876:Hughmilleria socialis
872:
683:In 1856, the species
627:Fossils described by
626:
575:(head) and posterior
2597:World Spider Catalog
2571:Novojilov, N. 1962:
1654:Erettopterus bilobus
1603:Acutiramus bohemicus
685:Pterygotus acuminata
436:(the "head" plate).
2690:2007PPP...252..557T
2631:2013GondR..23..354L
2383:2006GeoM..143..723T
2371:Geological Magazine
2293:1988Letha..21...13P
2239:1989JPal...63..642T
2050:2019JVPal..39E0247O
1992:2009Palgy..52.1141T
1538:Pterygotus anglicus
749:but did not assign
731:In 1903, the genus
689:John William Salter
660:, a Swiss-American
629:John William Salter
619:History of research
248:Pterygotus anglicus
2965:Onychopterelloidea
1480:Slimonia acuminata
1382:Hughmilleria wangi
1313:
1281:Internal phylogeny
888:External phylogeny
885:
873:Reconstruction of
753:to any particular
646:
512:Hughmilleria wangi
478:(top) and derived
396:Hughmilleria wangi
123:Slimonia acuminata
3528:
3527:
3466:Taxon identifiers
3457:
3456:
3138:
3137:
3134:
3133:
3130:
3129:
3086:Adelophthalmoidea
3068:Waeringopteroidea
3040:Carcinosomatoidea
2973:Onychopterellidae
2948:
2947:
2619:Gondwana Research
2573:Order Eurypterida
1930:Adelophthalmoidea
1802:
1801:
1793:
1792:
1784:
1783:
1775:
1774:
1766:
1765:
1757:
1756:
1748:
1747:
1739:
1738:
1730:
1729:
1632:
1631:
1623:
1622:
1614:
1613:
1434:
1433:
1277:
1276:
1268:
1267:
1259:
1258:
1250:
1249:
1241:
1240:
1232:
1231:
1223:
1222:
1214:
1213:
1205:
1204:
1196:
1195:
1187:
1186:
1080:Adelophthalmoidea
1058:Waeringopteroidea
1036:Carcinosomatoidea
965:
964:
914:genital operculum
906:Adelophthalmoidea
902:Carcinosomatoidea
762:John Mason Clarke
674:Old Red Sandstone
631:as belonging to "
321:
320:
315:
309:Hughmillerioidea
306:
233:
3563:
3521:
3520:
3508:
3507:
3506:
3493:
3492:
3491:
3461:
3460:
3449:
3448:
3423:Related articles
3380:Merostomichnites
3146:
3094:Adelophthalmidae
3076:Waeringopteridae
3048:Carcinosomatidae
3036:
3035:
3019:Strobilopteridae
2961:
2960:
2935:Hibbertopteridae
2884:Parastylonuridae
2854:
2853:
2827:
2826:
2738:
2731:
2724:
2715:
2714:
2709:
2708:
2706:
2700:. Archived from
2684:(3–4): 557–574.
2673:
2664:
2643:
2642:
2610:
2601:
2600:
2594:
2585:
2576:
2569:
2563:
2562:
2522:
2513:
2512:
2496:
2490:
2489:
2449:
2443:
2442:
2422:
2411:
2410:
2366:
2349:
2343:
2332:
2326:
2319:
2313:
2312:
2276:
2267:
2266:
2222:
2216:
2215:
2205:
2194:
2193:
2183:
2149:
2143:
2142:
2128:
2086:
2080:
2079:
2069:
2026:
2020:
2019:
1986:(5): 1141–1148.
1975:
1918:Llandovery epoch
1694:
1693:
1670:
1669:
1646:
1645:
1578:
1577:
1554:
1553:
1530:
1529:
1523:
1522:
1499:
1498:
1472:
1471:
1448:
1447:
1398:
1397:
1374:
1373:
1364:
1363:
1170:
1169:
1146:
1145:
1122:
1121:
1098:
1097:
1073:
1072:
1051:
1050:
1029:
1028:
1004:
1003:
989:Megalograptoidea
982:
981:
948:
947:
941:
940:
931:
930:
910:waeringopteroids
766:Rudolf Ruedemann
745:and the derived
739:Vernon Formation
469:Restorations of
463:
452:
310:
301:
224:
219:
206:
193:
143:
142:
115:
105:
42:
27:Temporal range:
21:
20:
3571:
3570:
3566:
3565:
3564:
3562:
3561:
3560:
3531:
3530:
3529:
3524:
3516:
3511:
3502:
3501:
3496:
3487:
3486:
3481:
3468:
3458:
3453:
3439:
3418:
3409:Chasmataspidida
3392:
3358:
3255:Campylocephalus
3228:
3185:
3147:
3126:
3112:Hughmilleriidae
3098:
3080:
3062:
3053:Megalograptidae
3029:
3023:
3009:Dolichopteridae
2995:
2983:Moselopteroidea
2977:
2954:
2944:
2930:Drepanopteridae
2916:
2912:Hardieopteridae
2899:Kokomopteroidea
2893:
2870:
2847:
2841:
2818:
2747:
2742:
2712:
2704:
2671:
2665:
2646:
2611:
2604:
2592:
2586:
2579:
2570:
2566:
2523:
2516:
2497:
2493:
2450:
2446:
2423:
2414:
2367:
2352:
2333:
2329:
2320:
2316:
2277:
2270:
2223:
2219:
2206:
2197:
2159:Biology Letters
2150:
2146:
2096:Biology Letters
2087:
2083:
2044:(3): e1620247.
2027:
2023:
1976:
1963:
1959:
1942:
1916:, are from the
1808:
1803:
1794:
1785:
1776:
1767:
1758:
1749:
1740:
1731:
1633:
1624:
1615:
1435:
1356:in a number of
1345:Herefordopterus
1283:
1278:
1269:
1260:
1251:
1242:
1233:
1224:
1215:
1206:
1197:
1188:
1130:Herefordopterus
966:
890:
867:
621:
562:Hughmilleriidae
528:Like all other
491:
490:
489:
488:
466:
465:
464:
455:
454:
453:
442:
406:supercontinents
382:and the famous
368:adelophthalmoid
355:members of the
271:Hughmilleriidae
257:
251:
223:
217:
204:
191:
137:
106:
104:
103:
98:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
38:442–372.2
37:
36:
25:
17:
12:
11:
5:
3569:
3559:
3558:
3553:
3548:
3543:
3541:Pterygotioidea
3526:
3525:
3523:
3522:
3509:
3494:
3478:
3476:
3474:Pterygotioidea
3470:
3469:
3455:
3454:
3444:
3441:
3440:
3438:
3437:
3432:
3426:
3424:
3420:
3419:
3417:
3416:
3411:
3406:
3400:
3398:
3397:Related groups
3394:
3393:
3391:
3390:
3383:
3376:
3368:
3366:
3360:
3359:
3357:
3356:
3349:
3342:
3335:
3328:
3325:Onychopterella
3321:
3314:
3307:
3300:
3293:
3286:
3283:Hibbertopterus
3279:
3272:
3265:
3258:
3251:
3244:
3241:Adelophthalmus
3236:
3234:
3233:Notable genera
3230:
3229:
3227:
3226:
3221:
3216:
3211:
3206:
3201:
3195:
3193:
3187:
3186:
3184:
3183:
3178:
3173:
3168:
3163:
3157:
3155:
3149:
3148:
3141:
3139:
3136:
3135:
3132:
3131:
3128:
3127:
3125:
3124:
3119:
3114:
3108:
3106:
3104:Pterygotioidea
3100:
3099:
3097:
3096:
3090:
3088:
3082:
3081:
3079:
3078:
3072:
3070:
3064:
3063:
3061:
3060:
3055:
3050:
3044:
3042:
3033:
3031:Diploperculata
3025:
3024:
3022:
3021:
3016:
3011:
3005:
3003:
2997:
2996:
2994:
2993:
2991:Moselopteridae
2987:
2985:
2979:
2978:
2976:
2975:
2969:
2967:
2958:
2950:
2949:
2946:
2945:
2943:
2942:
2937:
2932:
2926:
2924:
2918:
2917:
2915:
2914:
2909:
2907:Kokomopteridae
2903:
2901:
2895:
2894:
2892:
2891:
2886:
2880:
2878:
2872:
2871:
2869:
2868:
2862:
2860:
2858:Rhenopteroidea
2851:
2843:
2842:
2840:
2839:
2833:
2831:
2824:
2820:
2819:
2817:
2816:
2807:
2798:
2789:
2780:
2771:
2765:
2759:
2752:
2749:
2748:
2741:
2740:
2733:
2726:
2718:
2711:
2710:
2707:on 2011-07-18.
2644:
2625:(1): 354–366.
2602:
2577:
2564:
2537:(4): 725–736.
2514:
2491:
2444:
2433:(2): 331–361.
2412:
2377:(5): 723–735.
2350:
2327:
2314:
2268:
2233:(5): 642–657.
2217:
2195:
2166:(1): 106–109.
2144:
2103:(2): 265–269.
2081:
2021:
1960:
1958:
1955:
1954:
1953:
1948:
1941:
1938:
1816:Eastern Europe
1807:
1804:
1800:
1799:
1796:
1795:
1791:
1790:
1787:
1786:
1782:
1781:
1778:
1777:
1773:
1772:
1769:
1768:
1764:
1763:
1760:
1759:
1755:
1754:
1751:
1750:
1746:
1745:
1742:
1741:
1737:
1736:
1733:
1732:
1728:
1727:
1724:
1723:
1714:
1711:
1710:
1707:
1706:
1697:
1692:
1690:
1687:
1686:
1683:
1682:
1673:
1668:
1666:
1663:
1662:
1659:
1658:
1649:
1644:
1642:
1639:
1638:
1635:
1634:
1630:
1629:
1626:
1625:
1621:
1620:
1617:
1616:
1612:
1611:
1608:
1607:
1598:
1595:
1594:
1591:
1590:
1581:
1576:
1574:
1571:
1570:
1567:
1566:
1557:
1552:
1550:
1547:
1546:
1543:
1542:
1533:
1528:
1526:
1521:
1519:
1516:
1515:
1512:
1511:
1502:
1497:
1495:
1489:
1488:
1485:
1484:
1475:
1470:
1468:
1465:
1464:
1461:
1460:
1451:
1446:
1444:
1441:
1440:
1437:
1436:
1432:
1431:
1428:
1427:
1418:
1415:
1414:
1411:
1410:
1401:
1396:
1394:
1391:
1390:
1387:
1386:
1377:
1372:
1370:
1368:Pterygotioidea
1362:
1282:
1279:
1275:
1274:
1271:
1270:
1266:
1265:
1262:
1261:
1257:
1256:
1253:
1252:
1248:
1247:
1244:
1243:
1239:
1238:
1235:
1234:
1230:
1229:
1226:
1225:
1221:
1220:
1217:
1216:
1212:
1211:
1208:
1207:
1203:
1202:
1199:
1198:
1194:
1193:
1190:
1189:
1185:
1184:
1181:
1180:
1173:
1168:
1166:
1163:
1162:
1159:
1158:
1149:
1144:
1142:
1139:
1138:
1135:
1134:
1125:
1120:
1118:
1115:
1114:
1111:
1110:
1101:
1096:
1094:
1092:Pterygotioidea
1088:
1087:
1084:
1083:
1076:
1071:
1069:
1066:
1065:
1062:
1061:
1054:
1049:
1047:
1044:
1043:
1040:
1039:
1032:
1027:
1025:
1023:Diploperculata
1019:
1018:
1015:
1014:
1007:
1002:
1000:
997:
996:
993:
992:
985:
980:
978:
972:
971:
968:
967:
963:
962:
959:
958:
951:
946:
944:
939:
937:
929:
894:Diploperculata
889:
886:
866:
865:Classification
863:
620:
617:
495:Early Silurian
468:
467:
458:
457:
456:
447:
446:
445:
444:
443:
441:
438:
414:Early Silurian
376:hughmilleriids
360:Diploperculata
351:were the most
324:Pterygotioidea
319:
318:
317:
316:
307:
295:
294:
288:
287:
286:
285:
279:
273:
264:
263:
259:
258:
252:
242:
241:
235:
234:
221:Pterygotioidea
215:
211:
210:
208:Diploperculata
202:
198:
197:
189:
185:
184:
179:
175:
174:
169:
165:
164:
159:
155:
154:
149:
145:
144:
131:
130:
126:housed at the
117:
116:
108:
107:
99:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
29:Early Silurian
26:
24:Pterygotioidea
15:
9:
6:
4:
3:
2:
3568:
3557:
3554:
3552:
3549:
3547:
3544:
3542:
3539:
3538:
3536:
3519:
3514:
3510:
3505:
3504:Pterygotoidea
3499:
3495:
3490:
3484:
3480:
3479:
3477:
3475:
3471:
3467:
3462:
3452:
3442:
3436:
3433:
3431:
3428:
3427:
3425:
3421:
3415:
3412:
3410:
3407:
3405:
3402:
3401:
3399:
3395:
3389:
3388:
3384:
3382:
3381:
3377:
3375:
3374:
3370:
3369:
3367:
3365:
3361:
3355:
3354:
3350:
3348:
3347:
3343:
3341:
3340:
3336:
3334:
3333:
3332:Pentecopterus
3329:
3327:
3326:
3322:
3320:
3319:
3315:
3313:
3312:
3308:
3306:
3305:
3304:Megalograptus
3301:
3299:
3298:
3297:Jaekelopterus
3294:
3292:
3291:
3287:
3285:
3284:
3280:
3278:
3277:
3273:
3271:
3270:
3269:Drepanopterus
3266:
3264:
3263:
3259:
3257:
3256:
3252:
3250:
3249:
3248:Brachyopterus
3245:
3243:
3242:
3238:
3237:
3235:
3231:
3225:
3224:South America
3222:
3220:
3217:
3215:
3214:North America
3212:
3210:
3207:
3205:
3202:
3200:
3197:
3196:
3194:
3192:
3188:
3182:
3179:
3177:
3176:Carboniferous
3174:
3172:
3169:
3167:
3164:
3162:
3159:
3158:
3156:
3154:
3153:Geochronology
3150:
3145:
3123:
3120:
3118:
3115:
3113:
3110:
3109:
3107:
3105:
3101:
3095:
3092:
3091:
3089:
3087:
3083:
3077:
3074:
3073:
3071:
3069:
3065:
3059:
3056:
3054:
3051:
3049:
3046:
3045:
3043:
3041:
3037:
3034:
3032:
3026:
3020:
3017:
3015:
3012:
3010:
3007:
3006:
3004:
3002:
3001:Eurypteroidea
2998:
2992:
2989:
2988:
2986:
2984:
2980:
2974:
2971:
2970:
2968:
2966:
2962:
2959:
2957:
2951:
2941:
2940:Mycteroptidae
2938:
2936:
2933:
2931:
2928:
2927:
2925:
2923:
2922:Mycteropoidea
2919:
2913:
2910:
2908:
2905:
2904:
2902:
2900:
2896:
2890:
2887:
2885:
2882:
2881:
2879:
2877:
2876:Stylonuroidea
2873:
2867:
2866:Rhenopteridae
2864:
2863:
2861:
2859:
2855:
2852:
2850:
2844:
2838:
2835:
2834:
2832:
2828:
2825:
2821:
2815:
2814:Sclerophorata
2811:
2808:
2806:
2802:
2799:
2797:
2793:
2790:
2788:
2784:
2781:
2779:
2778:Euchelicerata
2775:
2772:
2770:
2766:
2764:
2760:
2758:
2754:
2753:
2750:
2746:
2739:
2734:
2732:
2727:
2725:
2720:
2719:
2716:
2703:
2699:
2695:
2691:
2687:
2683:
2679:
2678:
2670:
2663:
2661:
2659:
2657:
2655:
2653:
2651:
2649:
2640:
2636:
2632:
2628:
2624:
2620:
2616:
2609:
2607:
2598:
2591:
2584:
2582:
2574:
2568:
2560:
2556:
2552:
2548:
2544:
2540:
2536:
2532:
2528:
2521:
2519:
2510:
2506:
2502:
2495:
2487:
2483:
2479:
2475:
2471:
2467:
2463:
2459:
2455:
2448:
2440:
2436:
2432:
2428:
2421:
2419:
2417:
2408:
2404:
2400:
2396:
2392:
2388:
2384:
2380:
2376:
2372:
2365:
2363:
2361:
2359:
2357:
2355:
2347:
2341:
2340:
2331:
2324:
2318:
2310:
2306:
2302:
2298:
2294:
2290:
2286:
2282:
2275:
2273:
2264:
2260:
2256:
2252:
2248:
2244:
2240:
2236:
2232:
2228:
2221:
2214:. p. 23.
2213:
2212:
2204:
2202:
2200:
2191:
2187:
2182:
2177:
2173:
2169:
2165:
2161:
2160:
2155:
2148:
2140:
2136:
2132:
2127:
2122:
2118:
2114:
2110:
2106:
2102:
2098:
2097:
2092:
2085:
2077:
2073:
2068:
2063:
2059:
2055:
2051:
2047:
2043:
2039:
2038:
2033:
2025:
2017:
2013:
2009:
2005:
2001:
1997:
1993:
1989:
1985:
1981:
1980:Palaeontology
1974:
1972:
1970:
1968:
1966:
1961:
1952:
1949:
1947:
1944:
1943:
1937:
1933:
1931:
1927:
1923:
1919:
1915:
1910:
1908:
1904:
1900:
1899:South America
1896:
1892:
1889:(composed of
1888:
1884:
1880:
1876:
1875:South America
1872:
1868:
1864:
1860:
1856:
1852:
1848:
1843:
1841:
1837:
1833:
1829:
1825:
1824:North America
1821:
1817:
1813:
1798:
1797:
1789:
1788:
1780:
1779:
1771:
1770:
1762:
1761:
1753:
1752:
1744:
1743:
1735:
1734:
1726:
1725:
1722:
1721:
1720:
1713:
1712:
1709:
1708:
1705:
1704:
1703:
1696:
1695:
1689:
1688:
1685:
1684:
1681:
1680:
1679:
1672:
1671:
1665:
1664:
1661:
1660:
1657:
1656:
1655:
1648:
1647:
1641:
1640:
1637:
1636:
1628:
1627:
1619:
1618:
1610:
1609:
1606:
1605:
1604:
1597:
1596:
1593:
1592:
1589:
1588:
1587:
1580:
1579:
1573:
1572:
1569:
1568:
1565:
1564:
1563:
1556:
1555:
1549:
1548:
1545:
1544:
1541:
1540:
1539:
1532:
1531:
1525:
1524:
1518:
1517:
1514:
1513:
1510:
1509:
1508:
1501:
1500:
1494:
1491:
1490:
1487:
1486:
1483:
1482:
1481:
1474:
1473:
1467:
1466:
1463:
1462:
1459:
1458:
1457:
1450:
1449:
1443:
1442:
1439:
1438:
1430:
1429:
1426:
1425:
1424:
1417:
1416:
1413:
1412:
1409:
1408:
1407:
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1399:
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1383:
1376:
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1369:
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1355:
1349:
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1200:
1192:
1191:
1183:
1182:
1179:
1178:
1172:
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1165:
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1157:
1156:
1155:
1148:
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1133:
1132:
1131:
1124:
1123:
1117:
1116:
1113:
1112:
1109:
1108:
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1099:
1093:
1090:
1089:
1086:
1085:
1082:
1081:
1075:
1074:
1068:
1067:
1064:
1063:
1060:
1059:
1053:
1052:
1046:
1045:
1042:
1041:
1038:
1037:
1031:
1030:
1024:
1021:
1020:
1017:
1016:
1013:
1012:
1011:Eurypteroidea
1006:
1005:
999:
998:
995:
994:
991:
990:
984:
983:
977:
974:
973:
970:
969:
961:
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956:
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949:
943:
942:
936:
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928:
926:
921:
917:
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911:
907:
903:
899:
895:
882:
878:
877:
871:
862:
860:
856:
852:
848:
844:
839:
837:
833:
832:
827:
823:
819:
815:
814:
809:
805:
801:
797:
793:
789:
785:
781:
780:
775:
771:
767:
763:
758:
756:
752:
748:
744:
740:
736:
735:
729:
727:
723:
719:
718:
713:
709:
706:
703:(such as the
702:
698:
694:
690:
687:was named by
686:
681:
679:
675:
671:
667:
663:
659:
658:Louis Agassiz
655:
651:
643:
642:
637:
636:problematicus
635:
630:
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616:
614:
610:
605:
603:
602:
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589:
584:
582:
578:
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570:
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547:
543:
539:
535:
531:
526:
524:
523:
518:
514:
513:
508:
504:
500:
499:Late Devonian
496:
486:
485:
481:
477:
476:
472:
462:
451:
437:
435:
429:
427:
423:
419:
415:
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389:
385:
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350:
349:Pterygotioids
346:
342:
338:
334:
333:
329:
325:
313:
308:
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300:Slimonioidea
299:
298:
296:
293:
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278:
274:
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268:
267:
265:
260:
255:
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216:
214:Superfamily:
213:
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209:
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199:
196:
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180:
177:
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136:
132:
129:
125:
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118:
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109:
102:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
41:
34:
33:Late Devonian
30:
22:
19:
3473:
3385:
3378:
3371:
3351:
3344:
3337:
3330:
3323:
3316:
3309:
3302:
3295:
3290:Hughmilleria
3288:
3281:
3274:
3267:
3260:
3253:
3246:
3239:
3122:Pterygotidae
3103:
3058:Mixopteridae
3014:Eurypteridae
2889:Stylonuridae
2809:
2800:
2791:
2782:
2773:
2702:the original
2681:
2675:
2622:
2618:
2596:
2572:
2567:
2534:
2530:
2511:: 1080–1108.
2508:
2504:
2494:
2464:(1): 15–18.
2461:
2457:
2447:
2430:
2426:
2374:
2370:
2345:
2337:
2330:
2317:
2287:(1): 13–27.
2284:
2280:
2230:
2226:
2220:
2210:
2163:
2157:
2147:
2100:
2094:
2084:
2041:
2035:
2024:
1983:
1979:
1934:
1914:Hughmilleria
1913:
1911:
1863:Saudi Arabia
1844:
1809:
1806:Distribution
1717:
1716:
1700:
1699:
1676:
1675:
1652:
1651:
1601:
1600:
1584:
1583:
1560:
1559:
1536:
1535:
1505:
1504:
1493:Pterygotidae
1478:
1477:
1454:
1453:
1421:
1420:
1404:
1403:
1380:
1379:
1367:
1354:paraphyletic
1350:
1343:
1339:Hughmilleria
1338:
1335:Hughmilleria
1334:
1332:
1328:sister-group
1323:
1319:Ciurcopterus
1317:
1314:
1308:Hughmilleria
1306:
1300:
1294:
1177:Pterygotidae
1175:
1152:
1151:
1128:
1127:
1106:Hughmilleria
1104:
1103:
1091:
1078:
1056:
1034:
1009:
987:
953:
924:
922:
918:
891:
881:Hughmilleria
880:
874:
858:
843:Leif Størmer
840:
835:
831:Grossopterus
829:
825:
822:Hughmilleria
821:
817:
813:Erettopterus
811:
808:Erettopterus
807:
803:
799:
795:
787:
784:Hughmilleria
783:
777:
773:
769:
759:
751:Hughmilleria
750:
746:
742:
734:Hughmilleria
732:
730:
725:
721:
715:
707:
705:type species
700:
684:
682:
669:
653:
647:
639:
632:
606:
599:
585:
566:
550:Pterygotidae
540:composed of
530:chelicerates
527:
520:
510:
492:
482:
473:
430:
402:
394:
392:
364:sister group
348:
330:
323:
322:
283:Pterygotidae
246:
239:Type species
220:
201:Infraorder:
121:
18:
3498:Wikispecies
3387:Palmichnium
3364:Ichnogenera
3262:Carcinosoma
2956:Eurypterina
2849:Stylonurina
2787:Prosomapoda
2769:Chelicerata
2767:Subphylum:
2745:Eurypterids
2067:10784/26939
1926:South China
1812:Scandinavia
1291:type genera
976:Eurypterina
955:Stylonurina
935:Eurypterida
898:Eurypterina
855:Eurypterina
847:superfamily
708:P. anglicus
641:nomen vanum
577:opisthosoma
554:exoskeleton
503:eurypterine
440:Description
422:South China
384:pterygotids
341:eurypterids
337:superfamily
195:Eurypterida
182:Chelicerata
178:Subphylum:
3535:Categories
3353:Stylonurus
3339:Pterygotus
3318:Mixopterus
3311:Megarachne
3276:Eurypterus
3161:Ordovician
3117:Slimonidae
3028:Infraorder
2805:Dekatriata
2796:Planaterga
2763:Arthropoda
2346:Pterygotus
2339:Formations
1957:References
1907:Antarctica
1296:Pterygotus
836:Pterygotus
804:Pterygotus
800:Curviramus
796:Pterygotus
788:Pterygotus
770:Pterygotus
747:Pterygotus
743:Eurypterus
726:Pterygotus
722:Pterygotus
712:David Page
701:Pterygotus
693:Lesmahagow
670:Pterygotus
654:Pterygotus
650:type genus
634:Pterygotus
609:chelicerae
601:Eurypterus
558:Slimonidae
517:arthropods
484:Pterygotus
480:pterygotid
388:cheliceral
374:and small
357:infraorder
345:arthropods
332:Pterygotus
328:type genus
277:Slimonidae
172:Arthropoda
120:Fossil of
3430:Metastoma
3414:Xiphosura
3404:Arachnida
3191:Geography
2755:Kingdom:
2559:140668235
2551:0022-3360
2486:131539776
2478:0371-6260
2399:1469-5081
2309:1502-3931
2255:0022-3360
2117:1744-9561
2076:198237241
2008:1475-4983
1903:Australia
1847:Australia
1836:Laurentia
896:, in the
818:Hastimima
792:subgenera
779:Hastimima
666:geologist
662:biologist
615:(claws).
548:. In the
534:segmented
487:(bottom).
426:Laurentia
380:slimonids
303:Novojilov
262:Families
230:Ruedemann
158:Kingdom:
152:Eukaryota
3489:Q7257013
3483:Wikidata
3451:Category
3373:Arcuites
3346:Slimonia
3171:Devonian
3166:Silurian
2953:Suborder
2846:Suborder
2823:Taxonomy
2761:Phylum:
2757:Animalia
2407:83835591
2263:46953627
2190:18029297
2135:19828493
2016:84268058
1940:See also
1922:Scotland
1887:Gondwana
1832:Avalonia
1324:Slimonia
1302:Slimonia
1154:Slimonia
859:Slimonia
826:Slimonia
774:Slimonia
717:Slimonia
697:Scotland
678:Scotland
542:proteins
507:carapace
475:Slimonia
471:slimonid
434:carapace
418:Scotland
362:and the
292:Synonyms
168:Phylum:
162:Animalia
148:Domain:
3219:Oceania
3181:Permian
2686:Bibcode
2627:Bibcode
2439:1301554
2379:Bibcode
2289:Bibcode
2281:Lethaia
2235:Bibcode
2181:2412931
2126:2865068
2046:Bibcode
1988:Bibcode
1883:Siberia
1879:Bohemia
1859:Florida
1851:Morocco
1828:Baltica
1820:Britain
810:), but
588:derived
573:prosoma
569:tagmata
538:cuticle
497:to the
390:claws.
366:of the
353:derived
312:Størmer
254:Agassiz
188:Order:
3518:286977
3209:Europe
3199:Africa
2830:Genera
2557:
2549:
2484:
2476:
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2074:
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1891:Africa
927:2013.
925:et al.
802:) and
755:family
613:chelae
592:telson
546:chitin
314:, 1974
305:, 1962
256:, 1844
232:, 1912
228:&
226:Clarke
2810:Clade
2801:Clade
2792:Clade
2783:Clade
2774:Clade
2705:(PDF)
2672:(PDF)
2593:(PDF)
2555:S2CID
2482:S2CID
2435:JSTOR
2403:S2CID
2259:S2CID
2072:S2CID
2012:S2CID
1895:India
1871:Paris
1867:China
1855:Libya
596:lance
581:basal
372:basal
3204:Asia
2547:ISSN
2474:ISSN
2395:ISSN
2305:ISSN
2251:ISSN
2186:PMID
2131:PMID
2113:ISSN
2004:ISSN
1924:and
1905:and
1881:and
1822:and
1305:and
824:and
782:and
764:and
664:and
560:and
544:and
420:and
46:PreꞒ
2694:doi
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2297:doi
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2176:PMC
2168:doi
2121:PMC
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1996:doi
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676:of
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