102:
811:(Isady locality, Mutovino), Permian, Lopingian, Wuchiapingian/Changhsingian (Tatarian, Severodvinskian/Vyatkian), −259.0–254.0 Ma; Udmurtia (Chepanikha, Rossokha River Valley), Permian, Guadalupian, Wordian (Biarmian, Urzhumian), −268.0–265.0 Ma; Kemerovo Region (Sokolova, Kutsnetsk Basin), Permian, Lopingian, Wuchiapingian/Changhsingian (Tatarian, Severodvinian/Vyatkian), −259.0–252.3 Ma; Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
123:
709:(Letopala River and Soyana), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma; Udmurtia (Chepanikha, Rossokha River Valley), Permian, Guadalupian, Wordian (Biarmian, Urzhumian), −268.0–265.0 Ma: Tatarstan (Mendeleevsk District, Tikhie Gory), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
584:(Kargala), Permian, Guadalupian/Lopingian, Capitanian/Wuchiapingian (Tatarian, Sweverodvinian), −265.0–259.0 Ma; Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma. One undescribed species of this genus was recorded from Indian
516:– Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma; Kemerovo Region (Tykhta River, Sokolova, Kutsnetsk Basin), Permian, Lopingian, Wuchiapingian/Changhsingian (Tatarian, Severodvinian/Vyatkian), −259.0–252.3 Ma.
325:
as subfamilies
Permocupedinae and Taldycupedinae respectively. However, other studies have recovered Talycupedidae as more closely related to crown-group beetles than to Permocupedidae.
868:– Brazil (Rio Grande do Sul (Road Cut on Road BR 290 near Minas do Leao), Irati Formation, Permian, Kungurian, −279.5–272.5 Ma; considered indeterminate to genus by Kirejtshuk (2020)
1020:
Zhao, Xianye; Yu, Yilun; Clapham, Matthew E; Yan, Evgeny; Chen, Jun; Jarzembowski, Edmund A; Zhao, Xiangdong; Wang, Bo (2021-11-08). Perry, George H; Fikacek, Martin (eds.).
942:[Permian fossil insects from the Arkhangelsk District. Part II. Neuroptera, Megaloptera and Coleoptera with the description of two new beetles from Tikhie Gory]
1346:
638:(Houdong, Yinping Mountain, SW Chaohu City), Permian, Guadalupian/Lopingian, Capitanian/Wuchiapingian (Tatarian, Severodvinian/Vyatkian), −265.0–259.0 Ma.
881:– Russia: Orenburg Region (Novo-Aleksandrovka), Permian, Guadalupian/Lopingian, Capitanian/Wuchiapingian (Tatarian, Severodvinian), −265.0–259.0 Ma.
1333:
939:Пермские ископаемые насекомые Архангельского края. Часть II. Сетчатокрылые, вислокрылые и жуки, с приложением описания двух новых жуков из Тихих Гор
456:– Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma).
554:– Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
434:– Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
412:– Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
1415:
1425:
1081:"Taxonomic Review of Fossil Coleopterous Families (Insecta, Coleoptera). Suborder Archostemata: Superfamilies Coleopseoidea and Cupedoidea"
907:
Kirov Region (Kityak), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
1177:
Ponomarenko, A. G.; Yan, E. V.; Huang, Di-ying (2014). "New beetles (Coleoptera) from the terminal Middle
Permian of China".
267:
stem group beetles. They first appeared during the Early
Permian, and were one of the dominant groups of beetles during the
101:
580:(Kityak), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma;
122:
967:"First Triassic record of the beetle family Permocupedidae (Insecta: Coleoptera): a peculiar example of a Lazarus taxon"
381:(Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma
218:
1128:
Boudinot, Brendon Elias; Yan, Evgeny
Viktorovich; Prokop, Jakub; Luo, Xiao-Zhu; Beutel, Rolf Georg (2022-07-28).
1130:"Permian parallelisms: Reanalysis of †Tshekardocoleidae sheds light on the earliest evolution of the Coleoptera"
937:
1420:
1405:
1400:
672:), Lower Permian, Cisuralian, Artinskian/Kungurian (Cisuralian/Biarmian, Kungurian/Kazanian), −279.5–272.5 Ma.
1410:
1250:
904:
1372:
771:, Permian, Cisuralian, Artinskian/Kungurian (Cisuralian/Biarmian, Kungurian/Kazanian), −279.5–272.5 Ma.
585:
1320:
1186:
1092:
978:
1288:
786:
547:
342:
569:
509:
479:
405:
281:
1159:
1002:
874:
706:
239:
117:
1377:
896:
691:
653:
360:
627:
427:
330:
1359:
1307:
1151:
1110:
1061:
1043:
994:
764:
756:
1364:
1351:
449:
1228:
1194:
1141:
1100:
1051:
1033:
986:
491:
294:
249:
768:
581:
487:
378:
286:
264:
200:
187:
1190:
1096:
982:
1056:
1021:
808:
665:
368:
322:
1394:
1163:
1006:
966:
669:
317:
59:
1312:
577:
338:
1273:
1282:
1105:
1080:
298:
268:
34:
1198:
990:
311:
79:
44:
1233:
1216:
1155:
1114:
1047:
998:
272:
154:
134:
84:
1065:
1299:
1267:
301:(wood eating), which is presumed to be the ancestral ecology of beetles.
276:
74:
69:
54:
49:
39:
1038:
1338:
1146:
1129:
1022:"Early evolution of beetles regulated by the end-Permian deforestation"
290:
89:
64:
174:
164:
144:
1244:
635:
1325:
1217:"Permian Insects from Paraná Basin, South Brazil, IV Coleoptera"
309:
Kirejtshuk (2020) included the following genera in an expanded (
1248:
965:
Ponomarenko, Alexandr G.; Bashkuev, Alexei S. (December 2018).
345:, Permian, Cisuralian, Sakmarian/Artinskian, −290.1–279.5 Ma.
399:– considered indeterminate to genus by Kirejtshuk (2020)
948:
494:
Middle
Triassic, Olenekian/Anisian, −247.2–242.0 Ma.
683:
Shcherbakov, Makarkin, Aristov & Vasilenko, 2009
662:
Shcherbakov, Makarkin, Aristov & Vasilenko, 2009
1257:
767:(Road Cut on Road BR 290 near Minas do Leao),
8:
1210:
1208:
297:of Germany. They are thought to have been
1245:
100:
20:
1232:
1145:
1104:
1055:
1037:
1079:Kirejtshuk, Alexander G. (2020-02-17).
928:
275:, with only one species known from the
7:
352:Geertsema & van den Heever, 1996
335:Geertsema & van den Heever, 1996
645:Ponomarenko, Yan & Huang, 2014
632:Ponomarenko, Yan & Huang, 2014
14:
501:Ponomarenko & Bashkuev, 2018
484:Ponomarenko & Bashkuev, 2018
315:) family, with the traditional (
121:
109:Fossil and life restoration of
1:
847:Protocupoides plavilstshikovi
321:) Permocupedidae and related
1416:Prehistoric animals of China
486:– Germany: Gambach am Main (
1426:Prehistoric insect families
1106:10.3390/geosciences10020073
863:"Protocupoides" rohdendorfi
520:Ichthyocupes kuznetskiensis
1442:
950:(in Russian and English).
271:. They became rare in the
1199:10.1134/S0031030114010109
991:10.1007/s12542-018-0426-4
245:
238:
231:
226:
118:Scientific classification
116:
108:
99:
23:
16:Extinct family of beetles
1234:10.22456/1807-9806.21678
938:
608:Kaltanicupes kitjakensis
600:Kaltanicupes kargalensis
1179:Paleontological Journal
936:Martynov, A.V. (1933).
831:Protocupoides elongatus
823:Protocupoides carinatus
558:Kaltanocoleus pospelovi
394:"Archicupes" reichardti
885:Tatarocupes granulatus
588:of the Lower Permian.
1373:Paleobiology Database
1134:Systematic Entomology
855:Protocupoides sharovi
775:Pintolla ponomarenkoi
746:Permocupes sojanensis
713:Permocupes distinctus
676:Maricoleus valentinae
642:Linicupes yinpinensis
616:Kaltanicupes richteri
536:Ichthyocupes tyzhnovi
468:Eocupes lukjanovitshi
416:Cytocupoides longatus
111:Permocupes sojanensis
1215:Pinto, I.D. (1987).
498:Frankencupes ultimus
385:Archicupes jacobsoni
282:Frankencupes ultimus
1191:2014PalJ...48..191P
1097:2020Geosc..10...73K
1039:10.7554/eLife.72692
983:2018PalZ...92..587P
914:(Ponomarenko, 1963)
905:Belebeevo Formation
839:Protocupoides esini
737:Permocupes semenovi
707:Arkhangel’sk Region
668:(Cape, Novosilsky,
592:Kaltanicupes acutus
343:Whitehill Formation
1147:10.1111/syen.12562
858:(Rohdendorf, 1961)
826:(Rohdendorf, 1961)
818:(Rohdendorf, 1961)
815:Protocupoides acer
721:Permocupes grandis
531:(Rohdendorf, 1961)
528:Ichthyocupes skoki
460:Eocupes cellulosus
438:Cytocupes angustus
30:Cisuralian–Anisian
1388:
1387:
1360:Open Tree of Life
1251:Taxon identifiers
901:Ponomarenko, 1969
888:Ponomarenko, 2004
879:Ponomarenko, 2004
842:Ponomarenko, 2013
834:Ponomarenko, 2013
765:Rio Grande do Sul
749:Ponomarenko, 1963
732:Ponomarenko, 2013
724:Ponomarenko, 1963
611:Ponomarenko, 1963
603:Ponomarenko, 1963
595:Ponomarenko, 1963
586:Barakar Formation
523:Ponomarenko, 1969
463:Ponomarenko, 1969
419:Ponomarenko, 1969
410:Ponomarenko, 1969
341:(Kenmoore Farm),
258:
257:
222:
1433:
1381:
1380:
1368:
1367:
1355:
1354:
1342:
1341:
1329:
1328:
1316:
1315:
1303:
1302:
1293:
1292:
1291:
1278:
1277:
1276:
1246:
1239:
1238:
1236:
1212:
1203:
1202:
1174:
1168:
1167:
1149:
1125:
1119:
1118:
1108:
1076:
1070:
1069:
1059:
1041:
1017:
1011:
1010:
962:
956:
955:
945:
933:
915:
911:Uralocupes major
902:
889:
880:
867:
859:
851:
850:Rohdendorf, 1961
843:
835:
827:
819:
806:
805:Rohdendorf, 1961
799:
798:Rohdendorf, 1961
792:
791:Rohdendorf, 1961
779:
762:
761:Kirejtshuk, 2020
750:
741:
733:
729:Permocupes latus
725:
717:
716:(Martynov, 1933)
704:
697:
684:
663:
646:
633:
620:
619:Rohdendorf, 1961
612:
604:
596:
575:
574:Rohdendorf, 1961
562:
561:Rohdendorf, 1961
553:
552:Rohdendorf, 1961
540:
539:Rohdendorf, 1961
532:
524:
515:
514:Rohdendorf, 1961
502:
485:
472:
471:Rohdendorf, 1961
464:
455:
454:Rohdendorf, 1961
442:
441:Rohdendorf, 1961
433:
432:Rohdendorf, 1961
420:
411:
398:
397:Rohdendorf, 1961
389:
388:Rohdendorf, 1961
376:
375:Rohdendorf, 1961
366:
365:Rohdendorf, 1961
353:
349:Afrocupes firmae
337:– South Africa:
336:
265:Protocoleopteran
253:
247:Kaltanocoleidae
217:
212:
199:
186:
126:
125:
104:
94:
31:
27:Temporal range:
21:
1441:
1440:
1436:
1435:
1434:
1432:
1431:
1430:
1421:Beetle families
1406:Permian insects
1401:Permocupedoidea
1391:
1390:
1389:
1384:
1376:
1371:
1363:
1358:
1350:
1345:
1337:
1332:
1324:
1319:
1311:
1306:
1298:
1296:
1287:
1286:
1281:
1272:
1271:
1266:
1253:
1243:
1242:
1214:
1213:
1206:
1176:
1175:
1171:
1127:
1126:
1122:
1078:
1077:
1073:
1019:
1018:
1014:
964:
963:
959:
943:
940:
935:
934:
930:
925:
913:
900:
887:
878:
865:
857:
849:
841:
833:
825:
817:
804:
797:
790:
777:
769:Irati Formation
760:
748:
739:
731:
723:
715:
702:
695:
678:
657:
644:
631:
618:
610:
602:
594:
582:Orenburg Region
573:
560:
551:
538:
530:
522:
513:
500:
488:Lower Franconia
483:
470:
462:
453:
440:
431:
418:
409:
396:
387:
379:Kemerovo Region
374:
364:
351:
334:
307:
287:Middle Triassic
263:is a family of
248:
216:
210:
201:Permocupedoidea
197:
188:Protocoleoptera
184:
120:
95:
93:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
42:
37:
29:
28:
25:
17:
12:
11:
5:
1439:
1437:
1429:
1428:
1423:
1418:
1413:
1408:
1403:
1393:
1392:
1386:
1385:
1383:
1382:
1369:
1356:
1343:
1330:
1317:
1304:
1294:
1289:Permocupedidae
1279:
1263:
1261:
1259:Permocupedidae
1255:
1254:
1249:
1241:
1240:
1204:
1185:(2): 191–200.
1169:
1120:
1071:
1012:
977:(4): 587–591.
957:
927:
926:
924:
921:
920:
919:
918:
917:
916:(Type species)
893:
892:
891:
890:(Type species)
871:
870:
869:
860:
852:
844:
836:
828:
820:
809:Vologda Region
783:
782:
781:
780:(Type species)
753:
752:
751:
743:
742:(Type species)
740:Martynov, 1933
734:
726:
718:
703:Martynov, 1933
696:Martynov, 1933
688:
687:
686:
685:(Type species)
666:Primorsky Krai
650:
649:
648:
647:(Type species)
624:
623:
622:
621:(Type species)
613:
605:
597:
566:
565:
564:
563:(Type species)
544:
543:
542:
541:(Type species)
533:
525:
506:
505:
504:
503:(Type species)
476:
475:
474:
473:(Type species)
465:
446:
445:
444:
443:(Type species)
424:
423:
422:
421:(Type species)
402:
401:
400:
391:
390:(Type species)
357:
356:
355:
354:(Type species)
323:Taldycupedidae
306:
303:
269:Middle Permian
261:Permocupedidae
256:
255:
243:
242:
236:
235:
229:
228:
224:
223:
214:Permocupedidae
208:
204:
203:
195:
191:
190:
182:
178:
177:
172:
168:
167:
162:
158:
157:
152:
148:
147:
142:
138:
137:
132:
128:
127:
114:
113:
106:
105:
97:
96:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
38:
33:
32:
26:
24:Permocupedidae
15:
13:
10:
9:
6:
4:
3:
2:
1438:
1427:
1424:
1422:
1419:
1417:
1414:
1412:
1411:Permian China
1409:
1407:
1404:
1402:
1399:
1398:
1396:
1379:
1374:
1370:
1366:
1361:
1357:
1353:
1348:
1344:
1340:
1335:
1331:
1327:
1322:
1318:
1314:
1309:
1305:
1301:
1295:
1290:
1284:
1280:
1275:
1269:
1265:
1264:
1262:
1260:
1256:
1252:
1247:
1235:
1230:
1226:
1222:
1218:
1211:
1209:
1205:
1200:
1196:
1192:
1188:
1184:
1180:
1173:
1170:
1165:
1161:
1157:
1153:
1148:
1143:
1139:
1135:
1131:
1124:
1121:
1116:
1112:
1107:
1102:
1098:
1094:
1090:
1086:
1082:
1075:
1072:
1067:
1063:
1058:
1053:
1049:
1045:
1040:
1035:
1031:
1027:
1023:
1016:
1013:
1008:
1004:
1000:
996:
992:
988:
984:
980:
976:
972:
968:
961:
958:
953:
949:
941:
932:
929:
922:
912:
909:
908:
906:
899:
898:
894:
886:
883:
882:
877:
876:
872:
864:
861:
856:
853:
848:
845:
840:
837:
832:
829:
824:
821:
816:
813:
812:
810:
803:
796:
789:
788:
787:Protocupoides
784:
778:(Pinto, 1987)
776:
773:
772:
770:
766:
759:
758:
754:
747:
744:
738:
735:
730:
727:
722:
719:
714:
711:
710:
708:
701:
700:Permocupoides
694:
693:
689:
682:
677:
674:
673:
671:
670:Russky Island
667:
661:
656:
655:
651:
643:
640:
639:
637:
630:
629:
625:
617:
614:
609:
606:
601:
598:
593:
590:
589:
587:
583:
579:
572:
571:
567:
559:
556:
555:
550:
549:
548:Kaltanocoleus
545:
537:
534:
529:
526:
521:
518:
517:
512:
511:
507:
499:
496:
495:
493:
492:Röt Formation
489:
482:
481:
477:
469:
466:
461:
458:
457:
452:
451:
447:
439:
436:
435:
430:
429:
425:
417:
414:
413:
408:
407:
403:
395:
392:
386:
383:
382:
380:
373:
370:
363:
362:
358:
350:
347:
346:
344:
340:
333:
332:
328:
327:
326:
324:
320:
319:
318:sensu stricto
314:
313:
304:
302:
300:
296:
295:Röt Formation
292:
288:
284:
283:
278:
274:
270:
266:
262:
254:
251:
244:
241:
237:
234:
230:
225:
220:
215:
209:
206:
205:
202:
196:
194:Superfamily:
193:
192:
189:
183:
180:
179:
176:
173:
170:
169:
166:
163:
160:
159:
156:
153:
150:
149:
146:
143:
140:
139:
136:
133:
130:
129:
124:
119:
115:
112:
107:
103:
98:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
41:
36:
22:
19:
1258:
1227:(19): 5–12.
1224:
1220:
1182:
1178:
1172:
1137:
1133:
1123:
1088:
1084:
1074:
1029:
1025:
1015:
974:
970:
960:
951:
947:
931:
910:
895:
884:
873:
862:
854:
846:
838:
830:
822:
814:
807:) – Russia:
801:
794:
785:
774:
755:
745:
736:
728:
720:
712:
705:) – Russia:
699:
690:
680:
679:Shcherbakov
675:
659:
658:Shcherbakov
652:
641:
626:
615:
607:
599:
591:
578:Kirov Region
570:Kaltanicupes
568:
557:
546:
535:
527:
519:
510:Ichthyocupes
508:
497:
480:Frankencupes
478:
467:
459:
448:
437:
426:
415:
406:Cytocupoides
404:
393:
384:
377:) – Russia,
371:
359:
348:
339:Western Cape
329:
316:
310:
308:
280:
273:Late Permian
260:
259:
246:
232:
213:
110:
18:
1283:Wikispecies
1085:Geosciences
875:Tatarocupes
866:Pinto, 1987
372:Palaeocupes
299:xylophagous
1395:Categories
1032:: e72692.
923:References
903:– Russia:
897:Uralocupes
795:Tomiocupes
763:– Brazil:
692:Permocupes
664:– Russia,
654:Maricoleus
576:– Russia:
361:Archicupes
312:sensu lato
250:Rohdendorf
181:Suborder:
175:Coleoptera
155:Arthropoda
1274:Q19385267
1221:Pesquisas
1164:251171914
1156:0307-6970
1140:: 69–96.
1115:2076-3263
1091:(2): 73.
1048:2050-084X
1007:133665417
999:0031-0220
634:– China:
628:Linicupes
428:Cytocupes
331:Afrocupes
285:from the
233:See text
141:Kingdom:
135:Eukaryota
1297:BioLib:
1268:Wikidata
1066:34747694
954:: 63–96.
802:Tricupes
757:Pintolla
305:Taxonomy
277:Triassic
240:Synonyms
219:Martynov
207:Family:
151:Phylum:
145:Animalia
131:Domain:
1365:4664797
1339:4769965
1326:8975518
1187:Bibcode
1093:Bibcode
1057:8585485
979:Bibcode
450:Eocupes
291:Anisian
227:Genera
171:Order:
165:Insecta
161:Class:
1378:153424
1352:103066
1300:854606
1162:
1154:
1113:
1064:
1054:
1046:
1005:
997:
944:(DjVu)
252:, 1961
221:, 1933
1347:IRMNG
1313:62679
1160:S2CID
1026:eLife
1003:S2CID
793:(syn
698:(syn
636:Anhui
1334:GBIF
1152:ISSN
1111:ISSN
1062:PMID
1044:ISSN
995:ISSN
971:PalZ
35:PreꞒ
1321:EoL
1308:CoL
1229:doi
1195:doi
1142:doi
1101:doi
1052:PMC
1034:doi
987:doi
490:),
369:syn
1397::
1375::
1362::
1349::
1336::
1323::
1310::
1285::
1270::
1225:19
1223:.
1219:.
1207:^
1193:.
1183:48
1181:.
1158:.
1150:.
1138:48
1136:.
1132:.
1109:.
1099:.
1089:10
1087:.
1083:.
1060:.
1050:.
1042:.
1030:10
1028:.
1024:.
1001:.
993:.
985:.
975:92
973:.
969:.
946:.
800:,
681:in
660:in
293:)
279:,
85:Pg
1237:.
1231::
1201:.
1197::
1189::
1166:.
1144::
1117:.
1103::
1095::
1068:.
1036::
1009:.
989::
981::
952:2
367:(
289:(
211:†
198:†
185:†
90:N
80:K
75:J
70:T
65:P
60:C
55:D
50:S
45:O
40:Ꞓ
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