102:
143:
120:
393:
around 252 million years ago. Fossil genera of the family are classified in the subfamily
Eodromeinae, while the two modern genera are placed in the subfamily Trachypachinae. The two subfamilies are distinguished by the shape and position of the
884:
Vasilikopoulos, Alexandros; Balke, Michael; Kukowka, Sandra; Pflug, James M.; Martin, Sebastian; Meusemann, Karen; Hendrich, Lars; Mayer, Christoph; Maddison, David R.; Niehuis, Oliver; Beutel, Rolf G.; Misof, Bernhard (October 2021).
355:, and as a family been placed in several different positions relative to other beetle families. Recent molecular phylogenies have recovered as the sister group to the clade Carabidae+
997:
Ponomarenko, A. G.; Volkov, A. N. (2013). "Ademosynoides asiaticus
Martynov, 1936, the earliest known member of an extant beetle family (Insecta, Coleoptera, Trachypachidae)".
887:"Phylogenomic analyses clarify the pattern of evolution of Adephaga (Coleoptera) and highlight phylogenetic artefacts due to model misspecification and excessive data trimming"
1055:"An extraordinarily preserved new genus and species of Trachypachidae (Coleoptera, Adephaga) from the Early Jurassic of Germany and a review of fossil trachypachid genera"
3194:
3220:
351:, "the most interesting thing about this small family is its uncertain phylogenetic relationships"; it has previously been classified as a subfamily of
3005:
3168:
3207:
1118:
3294:
101:
3212:
142:
942:
Baca, Stephen M.; Gustafson, Grey T.; Alexander, Alana M.; Gough, Harlan M.; Toussaint, Emmanuel F. A. (October 2021).
3090:
3147:
348:
3225:
3160:
3289:
1111:
459:
3020:
528:
445:
426:
3095:
610:
3246:
563:
623:
515:
3129:
3067:
2257:
1054:
1006:
955:
898:
819:
722:
606:
596:
Ponomarenko, 1977 (6+ species) Kazakhstan, Mongolia, Russia, China?, Middle
Jurassic-Early Cretaceous
466:
452:
110:
717:
301:
of their rearmost legs. There are only six known extant species in the family, with four species of
3058:
2476:
1104:
861:
618:
239:
730:
692:
674:
510:
365:
325:
and
Argentina. They were much more diverse in the past, with dozens of described species from the
1082:
1030:
979:
924:
683:
665:
592:
558:
540:
501:
137:
523:
484:
3251:
752:
656:
377:
3284:
3233:
3116:
2745:
2162:
1074:
1022:
971:
916:
865:
837:
774:
697:
601:
419:
398:
that are jointed to the mesocoxae (the section where the middle pair of legs joins the body).
371:
129:
3261:
3199:
639:
571:
3238:
2842:
2621:
2616:
2539:
2297:
1313:
1066:
1014:
963:
906:
870:
827:
580:
286:
2531:
2484:
2329:
2313:
1989:
1749:
1718:
1633:
1551:
1442:
1410:
489:
439:
943:
1010:
959:
902:
823:
3155:
3142:
3072:
2934:
2885:
2740:
2547:
2215:
2199:
2138:
2033:
2004:
1932:
1896:
1880:
1796:
1741:
1646:
1543:
1392:
1334:
1230:
857:
553:
Ponomarenko, 1977 (10+ species) China Russia
Mongolia, Middle Jurassic-Early Cretaceous
340:
forests, and are thought to be mostly predators and scavengers, active during the day (
332:
Their habits are similar to those of the ground beetles; they are usually found in the
298:
3278:
2916:
2720:
2711:
2452:
2423:
2415:
2407:
2381:
2356:
2252:
2239:
2178:
2096:
1999:
1994:
1875:
1809:
1780:
1764:
1728:
1698:
1651:
1605:
1572:
1488:
1480:
1451:
1289:
1086:
983:
928:
832:
807:
312:
294:
59:
1034:
687:
Wang, Zhang et
Ponomarenko, 2012, Yixian Formation, China, Early Cretaceous (Aptian)
3121:
2893:
2817:
2563:
2555:
2510:
2494:
2444:
2365:
2321:
2273:
2191:
2104:
1973:
1960:
1940:
1901:
1801:
1788:
1759:
1693:
1681:
1577:
1534:
1517:
1509:
1459:
1297:
1267:
1198:
1187:
1171:
435:
356:
303:
263:
1070:
363:
subgroup
Geadephaga. The earliest fossils known of this family are of the genera
3181:
3052:
2979:
2950:
2924:
2911:
2851:
2809:
2793:
2785:
2777:
2761:
2753:
2689:
2647:
2639:
2608:
2523:
2502:
2468:
2460:
2373:
2305:
2289:
2281:
2244:
2231:
2207:
2125:
2117:
2067:
2059:
2017:
1981:
1952:
1924:
1919:
1835:
1830:
1822:
1772:
1708:
1703:
1617:
1582:
1567:
1559:
1475:
1467:
1284:
411:
333:
317:
256:
119:
34:
2906:
2867:
2859:
2801:
2664:
2626:
2579:
2571:
2391:
2339:
2223:
2154:
2025:
2009:
1965:
1906:
1870:
1850:
1814:
1723:
1713:
1587:
1321:
1305:
1018:
341:
224:
214:
79:
44:
3043:
1096:
1078:
1026:
975:
920:
841:
756:
Kirejtshuk et
Ansorge, 2022, Kemerovo, Russia, Late Permian (Changhsingian)
678:
Ponomarenko, 1977 Kazakhstan, Mongolia, Middle/Late
Jurassic-Early Cretaceous
2971:
2963:
2955:
2942:
2898:
2697:
2672:
2600:
2428:
2399:
2083:
1914:
1888:
1840:
1754:
1656:
1625:
1597:
1430:
1405:
1400:
1381:
1346:
1235:
1218:
1210:
1159:
1147:
549:
532:
352:
174:
154:
84:
778:
Ponomarenko et Volkov, 2013 Kemerovo, Russia, Late
Permian (Changhsingian)
734:
Ponomarenko et Volkov, 2013 Kemerovo, Russia, Late Permian (Changhsingian)
3082:
3103:
3037:
2875:
2825:
2769:
2728:
2677:
2334:
2265:
2146:
1858:
1664:
1638:
1496:
1329:
1276:
1255:
1165:
648:
493:
395:
386:
360:
326:
204:
74:
69:
54:
49:
39:
3108:
944:"Integrative phylogenomics reveals a Permian origin of Adephaga beetles"
2592:
2584:
2436:
2170:
2091:
2075:
967:
911:
886:
701:
627:
382:
337:
308:
89:
64:
3186:
2183:
2043:
1676:
1153:
1141:
1128:
651:, China, Middle Jurassic, Glushkovo Formation, Russia, Upper Jurassic
631:
584:
390:
290:
194:
184:
164:
3173:
3014:
322:
3018:
1427:
1378:
1252:
1184:
1100:
3134:
669:
Bode, 1953 Posidonia Shale, Germany Early Jurassic (Toarcian)
806:
Wang, Bo; Zhang, Haichun; Ponomarenko, Alexandr G. (2012).
808:"Mesozoic Trachypachidae (Insecta: Coleoptera) from China"
1499:(furniture beetles, death watch beetles, spider beetles)
579:
Jia, Liang, Chang et Ren, 2011) (3 species) Mongolia,
1053:
Kirejtshuk, Alexander G.; Ansorge, Jörg (2022-05-17).
544:
Ponomarenko, 1985 (2 species) Russia, Upper Jurassic
3027:
2933:
2884:
2850:
2841:
2719:
2710:
2663:
2638:
2522:
2493:
2390:
2364:
2355:
2137:
2116:
2042:
1849:
1740:
1596:
1542:
1533:
1508:
1450:
1441:
1266:
647:Ponomarenko, 1977 preoccupied genus) (5 species)
2376:(jewel beetles, or metallic wood-boring beetles)
479:After revision in Kirejtshuk et Ansorge, 2022.
2099:(true weevils, bark beetles, ambrosia beetles)
1112:
8:
2342:(ironclad beetles, cylindrical bark beetles)
660:Ponomarenko, 1977 Kyrgyzstan, Lower Jurassic
3015:
2847:
2716:
2361:
1539:
1447:
1438:
1424:
1375:
1263:
1249:
1181:
1119:
1105:
1097:
505:Ren, 1995 China, Early Cretaceous (Aptian)
297:, but that are distinguished by the large
118:
100:
20:
3006:List of subgroups of the order Coleoptera
2086:(straight snout weevils, New York weevil)
910:
831:
2513:(cicada beetle, cicada parasite beetles)
798:
2603:(false fireflies, long-lipped beetles)
7:
3161:41d3291c-090b-4808-a887-2b65a62f731c
2629:(false metallic wood-boring beetles)
1048:
1046:
1044:
853:
851:
613:, Russia, Upper Jurassic (Tithonian)
566:, Germany Early Jurassic (Toarcian)
725:, China, Early Cretaceous (Aptian)
609:Kazakhstan, Middle-Late Jurassic,
14:
2240:Perimylopidae, or Promecheilidae
1825:(minute brown scavenger beetles)
833:10.1111/j.1475-4983.2012.01128.x
141:
2455:(variegated mud-loving beetles)
643:Kirejtshuk et Ansorge, 2022 (=
700:, Kyrgyzstan, Upper Triassic (
626:, Kazakhstan, Upper Triassic (
605:Ponomarenko, 1977 (2 species)
293:that generally resemble small
1:
2284:(narrow-waisted bark beetles)
1976:(parasitic flat bark beetles)
1927:(short-winged flower beetles)
1783:(lady beetles, or God's cows)
1520:(tooth-necked fungus beetles)
1071:10.1080/08912963.2022.2071709
2796:(sand-loving scarab beetles)
2186:(minute tree-fungus beetles)
2012:(silvanid flat bark beetles)
562:Kirejtshuk et Ansorge, 2022
2945:(primitive carrion beetles)
2901:(minute mud-loving beetles)
2764:(bumble bee scarab beetles)
2534:(soft-bodied plant beetles)
2505:(soft bodied plant beetles)
1308:(predaceous diving beetles)
583:, China, Early Cretaceous (
3311:
2780:(scavenger scarab beetles)
2772:(enigmatic scarab beetles)
1945:Lamingtonium binnaberrense
1659:(soft-wing flower beetles)
492:, Germany Early Jurassic (
347:According to entomologist
3003:
2919:(water scavenger beetles)
2652:Rhinorhipus tamborinensis
2431:(long-toed water beetles)
2324:(polypore fungus beetles)
2226:(palm and flower beetles)
2210:(tumbling flower beetles)
2157:(ant-like flower beetles)
1935:(lined flat bark beetles)
1909:(pleasing fungus beetles)
1804:(handsome fungus beetles)
1437:
1423:
1388:
1374:
1349:(burrowing water beetles)
1262:
1248:
1194:
1180:
1136:
1019:10.1134/S0031030113060063
874:(CRC Press, 2001), vol. 1
531:Mongolia, Late Jurassic (
407:Subfamily Trachypachinae
252:
247:
138:Scientific classification
136:
126:
117:
108:
99:
23:
3295:Beetles of North America
2966:(feather-winged beetles)
2300:(false longhorn beetles)
2247:(jugular-horned beetles)
2194:(false darkling beetles)
2165:(cryptic fungus beetles)
1984:(shining flower beetles)
1324:(crawling water beetles)
1223:Sikhotealinia zhiltzovae
460:Trachypachus zetterstedi
270:Fossil genera, see text
2447:(forest stream beetles)
2308:(synchroa bark beetles)
2268:(dead log bark beetles)
2260:(fire-coloured beetles)
2149:(ant-like leaf beetles)
1883:(silken fungus beetles)
1817:(well polished beetles)
1791:(minute fungus beetles)
1610:Acanthocnemus nigricans
999:Paleontological Journal
860:, "Trachypachidae", in
743:Ademosynoides asiaticus
518:, Russia, Late Jurassic
2958:(round fungus beetles)
2292:(false flower beetles)
2276:(wedge-shaped beetles)
2234:(false blister beetle)
2218:(hairy fungus beetles)
2173:(conifer bark beetles)
2070:(leaf-rolling weevils)
1731:(bark-gnawing beetles)
1491:(wounded-tree beetles)
1357:(false ground beetles)
1279:(trout-stream beetles)
739:Petrodromeus asiaticus
476:Subfamily †Eodromeinae
446:Trachypachus holmbergi
427:Systolosoma lateritium
3247:Paleobiology Database
2870:(false clown beetles)
2692:(plate-thigh beetles)
2574:(false click beetles)
2479:(water-penny beetles)
2128:(ship-timber beetles)
2107:(pine flower weevils)
1955:(root-eating beetles)
1775:(minute bark beetles)
1686:Phloiophilus edwardsi
1213:(reticulated beetles)
948:Systematic Entomology
891:Systematic Entomology
575:Ponomarenko, 1986 (=
315:, and two species of
30:Changhsingian–Present
3156:Fauna Europaea (new)
2748:(false stag beetles)
2733:Belohina inexpectata
2595:(net-winged beetles)
2558:(rare click beetles)
2471:(travertine beetles)
2463:(minute mud beetles)
2028:(dry-fungus beetles)
1863:Agapytho foveicollis
1620:(false skin beetles)
1483:(Jacobson's beetles)
1203:Crowsoniella relicta
723:Lushangfen Formation
607:Karabastau Formation
529:Ulaan-Ereg Formation
467:Trachypachus slevini
453:Trachypachus gibbsii
283:false ground beetles
281:(sometimes known as
111:Trachypachus gibbsii
2746:Diphyllostomatidae
2078:(primitive weevils)
1891:(flat bark beetles)
1641:(checkered beetles)
1628:(fruitworm beetles)
1316:(whirligig beetles)
1011:2013PalJ...47..601P
960:2021SysEn..46..968B
903:2021SysEn..46..991V
862:Ross H. Arnett, Jr.
824:2012Palgy..55..341W
611:Glushkovo Formation
2611:(glowworm beetles)
2316:(darkling beetles)
2163:Archeocrypticidae
2020:(palmetto beetles)
1767:(dry bark beetles)
1554:(longhorn beetles)
1059:Historical Biology
968:10.1111/syen.12506
912:10.1111/syen.12508
765:Petrodromeus minor
709:Sogdodromeus altus
696:Ponomarenko, 1977
622:Ponomarenko, 1977
527:Ponomarenko, 1989
514:Ponomarenko, 1985
307:found in northern
3272:
3271:
3234:Open Tree of Life
3021:Taxon identifiers
3012:
3011:
2999:
2998:
2995:
2994:
2991:
2990:
2983:
2975:
2974:(carrion beetles)
2967:
2959:
2946:
2920:
2902:
2871:
2863:
2837:
2836:
2829:
2821:
2813:
2805:
2797:
2789:
2781:
2773:
2765:
2757:
2749:
2736:
2706:
2705:
2693:
2685:
2655:
2630:
2612:
2604:
2596:
2588:
2575:
2567:
2559:
2551:
2550:(soldier beetles)
2543:
2540:Brachypsectridae
2535:
2514:
2506:
2480:
2472:
2464:
2456:
2448:
2440:
2432:
2419:
2411:
2403:
2377:
2351:
2350:
2343:
2325:
2317:
2309:
2301:
2298:Stenotrachelidae
2293:
2285:
2277:
2269:
2261:
2248:
2235:
2227:
2219:
2211:
2203:
2202:(blister beetles)
2195:
2187:
2174:
2166:
2158:
2150:
2129:
2108:
2100:
2087:
2079:
2071:
2063:
2053:
2029:
2021:
2013:
1985:
1977:
1969:
1956:
1948:
1936:
1928:
1910:
1892:
1884:
1866:
1826:
1818:
1805:
1792:
1784:
1776:
1768:
1750:Akalyptoischiidae
1732:
1689:
1672:
1660:
1642:
1629:
1621:
1613:
1563:
1555:
1529:
1528:
1521:
1500:
1492:
1484:
1471:
1463:
1419:
1418:
1396:
1370:
1369:
1366:
1365:
1358:
1350:
1342:
1325:
1317:
1309:
1301:
1293:
1280:
1244:
1243:
1226:
1214:
1206:
866:Michael C. Thomas
745:; Martynov, 1936)
698:Madygen Formation
624:Tologoi Formation
516:Ichetuy Formation
420:Systolosoma breve
415:Chile, Argentina
275:
274:
243:
130:Systolosoma breve
16:Family of beetles
3302:
3265:
3264:
3255:
3254:
3242:
3241:
3229:
3228:
3216:
3215:
3203:
3202:
3190:
3189:
3177:
3176:
3164:
3163:
3151:
3150:
3138:
3137:
3125:
3124:
3112:
3111:
3099:
3098:
3086:
3085:
3076:
3075:
3063:
3062:
3061:
3048:
3047:
3046:
3016:
2981:
2973:
2965:
2957:
2944:
2918:
2900:
2869:
2861:
2848:
2843:Staphyliniformia
2827:
2819:
2811:
2803:
2795:
2787:
2779:
2771:
2763:
2755:
2747:
2730:
2717:
2691:
2682:Declinia relicta
2679:
2649:
2628:
2622:Sinopyrophoridae
2617:Rhagophthalmidae
2610:
2602:
2594:
2586:
2573:
2565:
2557:
2549:
2541:
2533:
2532:Artematopodidae
2512:
2504:
2478:
2470:
2462:
2454:
2446:
2439:(riffle beetles)
2438:
2430:
2418:(turtle beetles)
2417:
2409:
2401:
2375:
2362:
2341:
2323:
2315:
2307:
2299:
2291:
2283:
2275:
2267:
2259:
2246:
2233:
2225:
2217:
2209:
2201:
2193:
2185:
2172:
2164:
2156:
2148:
2127:
2106:
2098:
2085:
2077:
2069:
2062:(fungus weevils)
2061:
2052:
2051:
2047:
2034:Tasmosalpingidae
2027:
2019:
2011:
1983:
1975:
1967:
1954:
1942:
1934:
1926:
1908:
1890:
1882:
1860:
1824:
1816:
1803:
1790:
1782:
1774:
1766:
1730:
1683:
1666:
1658:
1640:
1627:
1619:
1607:
1606:Acanthocnemidae
1561:
1553:
1540:
1519:
1498:
1490:
1482:
1469:
1461:
1448:
1439:
1425:
1394:
1376:
1356:
1348:
1336:
1323:
1315:
1307:
1299:
1292:(ground beetles)
1291:
1278:
1264:
1250:
1220:
1212:
1200:
1199:Crowsoniellidae
1182:
1121:
1114:
1107:
1098:
1091:
1090:
1050:
1039:
1038:
994:
988:
987:
939:
933:
932:
914:
881:
875:
871:American Beetles
855:
846:
845:
835:
803:
581:Yixian Formation
238:
146:
145:
122:
104:
94:
31:
27:Temporal range:
21:
3310:
3309:
3305:
3304:
3303:
3301:
3300:
3299:
3290:Beetle families
3275:
3274:
3273:
3268:
3260:
3258:
3250:
3245:
3237:
3232:
3224:
3219:
3211:
3206:
3198:
3193:
3185:
3180:
3172:
3167:
3159:
3154:
3146:
3141:
3133:
3128:
3120:
3115:
3107:
3102:
3094:
3089:
3081:
3079:
3071:
3066:
3057:
3056:
3051:
3042:
3041:
3036:
3023:
3013:
3008:
2987:
2929:
2880:
2862:(clown beetles)
2833:
2804:(betsy beetles)
2712:Scarabaeiformia
2702:
2659:
2634:
2566:(click beetles)
2542:(Texas beetles)
2518:
2489:
2485:Ptilodactylidae
2410:(cedar beetles)
2386:
2347:
2330:Trictenotomidae
2216:Mycetophagidae
2133:
2112:
2049:
2048:
2046:
2038:
1990:Phloeostichidae
1941:Lamingtoniidae
1933:Laemophloeidae
1881:Cryptophagidae
1845:
1736:
1719:Thanerocleridae
1682:Phloiophilidae
1669:Metaxina ornata
1634:Chaetosomatidae
1592:
1525:
1504:
1462:(auger beetles)
1443:Bostrichiformia
1433:
1415:
1411:Torridincolidae
1395:(skiff beetles)
1393:Hydroscaphidae
1384:
1362:
1355:Trachypachidae
1300:(tiger beetles)
1258:
1240:
1190:
1176:
1132:
1125:
1095:
1094:
1052:
1051:
1042:
996:
995:
991:
941:
940:
936:
897:(4): 991–1018.
883:
882:
878:
856:
849:
805:
804:
800:
795:
761:Apermunda minor
490:Posidonia Shale
404:
359:as part of the
237:
140:
95:
93:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
42:
37:
29:
28:
25:
17:
12:
11:
5:
3308:
3306:
3298:
3297:
3292:
3287:
3277:
3276:
3270:
3269:
3267:
3266:
3256:
3243:
3230:
3217:
3204:
3191:
3178:
3165:
3152:
3143:Fauna Europaea
3139:
3126:
3113:
3100:
3087:
3077:
3073:Trachypachidae
3064:
3059:Trachypachidae
3049:
3033:
3031:
3029:Trachypachidae
3025:
3024:
3019:
3010:
3009:
3004:
3001:
3000:
2997:
2996:
2993:
2992:
2989:
2988:
2986:
2985:
2982:(rove beetles)
2980:Staphylinidae
2977:
2969:
2961:
2953:
2948:
2939:
2937:
2935:Staphylinoidea
2931:
2930:
2928:
2927:
2922:
2917:Hydrophilidae
2914:
2909:
2904:
2896:
2890:
2888:
2886:Hydrophiloidea
2882:
2881:
2879:
2878:
2873:
2865:
2856:
2854:
2845:
2839:
2838:
2835:
2834:
2832:
2831:
2828:(hide beetles)
2823:
2815:
2812:(rain beetles)
2807:
2799:
2791:
2788:(stag beetles)
2783:
2775:
2767:
2759:
2751:
2743:
2741:Bolboceratidae
2738:
2725:
2723:
2714:
2708:
2707:
2704:
2703:
2701:
2700:
2695:
2687:
2675:
2669:
2667:
2661:
2660:
2658:
2657:
2648:Rhinorhipidae
2644:
2642:
2640:Rhinorhipoidea
2636:
2635:
2633:
2632:
2624:
2619:
2614:
2606:
2598:
2590:
2582:
2577:
2569:
2561:
2553:
2545:
2537:
2528:
2526:
2520:
2519:
2517:
2516:
2508:
2499:
2497:
2491:
2490:
2488:
2487:
2482:
2474:
2466:
2458:
2453:Heteroceridae
2450:
2442:
2434:
2426:
2421:
2416:Chelonariidae
2413:
2408:Callirhipidae
2405:
2402:(pill beetles)
2396:
2394:
2388:
2387:
2385:
2384:
2379:
2370:
2368:
2359:
2353:
2352:
2349:
2348:
2346:
2345:
2337:
2332:
2327:
2319:
2314:Tenebrionidae
2311:
2303:
2295:
2287:
2279:
2271:
2263:
2255:
2250:
2242:
2237:
2229:
2221:
2213:
2205:
2197:
2189:
2181:
2176:
2168:
2160:
2152:
2143:
2141:
2139:Tenebrionoidea
2135:
2134:
2132:
2131:
2122:
2120:
2114:
2113:
2111:
2110:
2102:
2097:Curculionidae
2094:
2089:
2081:
2073:
2065:
2056:
2054:
2044:Curculionoidea
2040:
2039:
2037:
2036:
2031:
2023:
2015:
2007:
2005:Protocucujidae
2002:
1997:
1992:
1987:
1979:
1971:
1963:
1958:
1950:
1938:
1930:
1922:
1917:
1912:
1904:
1899:
1897:Cybocephalidae
1894:
1886:
1878:
1873:
1868:
1855:
1853:
1847:
1846:
1844:
1843:
1838:
1833:
1828:
1820:
1812:
1807:
1799:
1797:Discolomatidae
1794:
1786:
1781:Coccinellidae
1778:
1770:
1765:Bothrideridae
1762:
1757:
1752:
1746:
1744:
1742:Coccinelloidea
1738:
1737:
1735:
1734:
1729:Trogossitidae
1726:
1721:
1716:
1711:
1706:
1701:
1696:
1691:
1679:
1674:
1662:
1654:
1649:
1647:Lophocateridae
1644:
1636:
1631:
1623:
1615:
1602:
1600:
1594:
1593:
1591:
1590:
1585:
1580:
1575:
1570:
1565:
1562:(leaf beetles)
1560:Chrysomelidae
1557:
1548:
1546:
1544:Chrysomeloidea
1537:
1531:
1530:
1527:
1526:
1524:
1523:
1514:
1512:
1506:
1505:
1503:
1502:
1494:
1489:Nosodendridae
1486:
1481:Jacobsoniidae
1478:
1473:
1470:(skin beetles)
1465:
1456:
1454:
1445:
1435:
1434:
1428:
1421:
1420:
1417:
1416:
1414:
1413:
1408:
1403:
1398:
1389:
1386:
1385:
1379:
1372:
1371:
1368:
1367:
1364:
1363:
1361:
1360:
1352:
1344:
1339:Meru phyllisae
1332:
1327:
1319:
1311:
1303:
1295:
1287:
1282:
1273:
1271:
1260:
1259:
1253:
1246:
1245:
1242:
1241:
1239:
1238:
1233:
1231:Micromalthidae
1228:
1216:
1208:
1195:
1192:
1191:
1185:
1178:
1177:
1175:
1174:
1168:
1162:
1156:
1150:
1144:
1137:
1134:
1133:
1126:
1124:
1123:
1116:
1109:
1101:
1093:
1092:
1065:(6): 958–975.
1040:
1005:(6): 601–606.
989:
954:(4): 968–990.
934:
876:
858:George E. Ball
847:
818:(2): 341–353.
797:
796:
794:
791:
790:
789:
788:
787:
786:
785:
770:
769:
768:
748:
747:
746:
726:
718:Xishanocarabus
713:
712:
711:
688:
679:
670:
661:
652:
635:
614:
597:
588:
567:
554:
545:
536:
519:
506:
497:
477:
474:
473:
472:
471:
470:
463:
456:
449:
432:
431:
430:
423:
403:
400:
295:ground beetles
279:Trachypachidae
273:
272:
268:
267:
260:
250:
249:
245:
244:
235:Trachypachidae
232:
228:
227:
222:
218:
217:
212:
208:
207:
202:
198:
197:
192:
188:
187:
182:
178:
177:
172:
168:
167:
162:
158:
157:
152:
148:
147:
134:
133:
124:
123:
115:
114:
106:
105:
97:
96:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
38:
33:
32:
26:
24:Trachypachidae
15:
13:
10:
9:
6:
4:
3:
2:
3307:
3296:
3293:
3291:
3288:
3286:
3283:
3282:
3280:
3263:
3257:
3253:
3248:
3244:
3240:
3235:
3231:
3227:
3222:
3218:
3214:
3209:
3205:
3201:
3196:
3192:
3188:
3183:
3179:
3175:
3170:
3166:
3162:
3157:
3153:
3149:
3144:
3140:
3136:
3131:
3127:
3123:
3118:
3114:
3110:
3105:
3101:
3097:
3092:
3088:
3084:
3078:
3074:
3069:
3065:
3060:
3054:
3050:
3045:
3039:
3035:
3034:
3032:
3030:
3026:
3022:
3017:
3007:
3002:
2984:
2978:
2976:
2970:
2968:
2962:
2960:
2954:
2952:
2949:
2947:
2941:
2940:
2938:
2936:
2932:
2926:
2923:
2921:
2915:
2913:
2910:
2908:
2905:
2903:
2897:
2895:
2892:
2891:
2889:
2887:
2883:
2877:
2874:
2872:
2868:Sphaeritidae
2866:
2864:
2858:
2857:
2855:
2853:
2849:
2846:
2844:
2840:
2830:
2824:
2822:
2818:Scarabaeidae
2816:
2814:
2808:
2806:
2800:
2798:
2792:
2790:
2784:
2782:
2776:
2774:
2768:
2766:
2760:
2758:
2756:(dor beetles)
2752:
2750:
2744:
2742:
2739:
2737:
2734:
2727:
2726:
2724:
2722:
2721:Scarabaeoidea
2718:
2715:
2713:
2709:
2699:
2696:
2694:
2688:
2686:
2683:
2676:
2674:
2671:
2670:
2668:
2666:
2662:
2656:
2653:
2646:
2645:
2643:
2641:
2637:
2631:
2625:
2623:
2620:
2618:
2615:
2613:
2607:
2605:
2599:
2597:
2591:
2589:
2583:
2581:
2578:
2576:
2570:
2568:
2562:
2560:
2556:Cerophytidae
2554:
2552:
2546:
2544:
2538:
2536:
2530:
2529:
2527:
2525:
2521:
2515:
2511:Rhipiceridae
2509:
2507:
2501:
2500:
2498:
2496:
2492:
2486:
2483:
2481:
2475:
2473:
2467:
2465:
2459:
2457:
2451:
2449:
2445:Eulichadidae
2443:
2441:
2435:
2433:
2427:
2425:
2424:Cneoglossidae
2422:
2420:
2414:
2412:
2406:
2404:
2398:
2397:
2395:
2393:
2389:
2383:
2382:Schizopodidae
2380:
2378:
2372:
2371:
2369:
2367:
2363:
2360:
2358:
2357:Elateriformia
2354:
2344:
2338:
2336:
2333:
2331:
2328:
2326:
2322:Tetratomidae
2320:
2318:
2312:
2310:
2304:
2302:
2296:
2294:
2288:
2286:
2280:
2278:
2274:Ripiphoridae
2272:
2270:
2264:
2262:
2258:Pyrochroidae
2256:
2254:
2253:Pterogeniidae
2251:
2249:
2243:
2241:
2238:
2236:
2230:
2228:
2222:
2220:
2214:
2212:
2206:
2204:
2198:
2196:
2192:Melandryidae
2190:
2188:
2182:
2180:
2179:Chalcodryidae
2177:
2175:
2169:
2167:
2161:
2159:
2153:
2151:
2145:
2144:
2142:
2140:
2136:
2130:
2124:
2123:
2121:
2119:
2115:
2109:
2105:Nemonychidae
2103:
2101:
2095:
2093:
2090:
2088:
2082:
2080:
2074:
2072:
2066:
2064:
2058:
2057:
2055:
2045:
2041:
2035:
2032:
2030:
2024:
2022:
2016:
2014:
2008:
2006:
2003:
2001:
2000:Propalticidae
1998:
1996:
1995:Priasilphidae
1993:
1991:
1988:
1986:
1980:
1978:
1974:Passandridae
1972:
1970:
1968:(sap beetles)
1964:
1962:
1959:
1957:
1951:
1949:
1946:
1939:
1937:
1931:
1929:
1923:
1921:
1918:
1916:
1913:
1911:
1905:
1903:
1900:
1898:
1895:
1893:
1887:
1885:
1879:
1877:
1876:Cavognathidae
1874:
1872:
1869:
1867:
1864:
1857:
1856:
1854:
1852:
1848:
1842:
1839:
1837:
1834:
1832:
1829:
1827:
1821:
1819:
1813:
1811:
1810:Eupsilobiidae
1808:
1806:
1802:Endomychidae
1800:
1798:
1795:
1793:
1789:Corylophidae
1787:
1785:
1779:
1777:
1771:
1769:
1763:
1761:
1758:
1756:
1753:
1751:
1748:
1747:
1745:
1743:
1739:
1733:
1727:
1725:
1722:
1720:
1717:
1715:
1712:
1710:
1707:
1705:
1704:Protopeltidae
1702:
1700:
1699:Prionoceridae
1697:
1695:
1692:
1690:
1687:
1680:
1678:
1675:
1673:
1670:
1663:
1661:
1655:
1653:
1652:Mauroniscidae
1650:
1648:
1645:
1643:
1637:
1635:
1632:
1630:
1624:
1622:
1616:
1614:
1611:
1604:
1603:
1601:
1599:
1595:
1589:
1586:
1584:
1581:
1579:
1576:
1574:
1573:Megalopodidae
1571:
1569:
1566:
1564:
1558:
1556:
1552:Cerambycidae
1550:
1549:
1547:
1545:
1541:
1538:
1536:
1532:
1522:
1518:Derodontidae
1516:
1515:
1513:
1511:
1510:Derodontoidea
1507:
1501:
1495:
1493:
1487:
1485:
1479:
1477:
1476:Endecatomidae
1474:
1472:
1466:
1464:
1460:Bostrichidae
1458:
1457:
1455:
1453:
1452:Bostrichoidea
1449:
1446:
1444:
1440:
1436:
1432:
1426:
1422:
1412:
1409:
1407:
1406:Sphaeriusidae
1404:
1402:
1399:
1397:
1391:
1390:
1387:
1383:
1377:
1373:
1359:
1353:
1351:
1345:
1343:
1340:
1333:
1331:
1328:
1326:
1320:
1318:
1312:
1310:
1304:
1302:
1298:Cicindelidae
1296:
1294:
1288:
1286:
1283:
1281:
1275:
1274:
1272:
1269:
1265:
1261:
1257:
1251:
1247:
1237:
1234:
1232:
1229:
1227:
1224:
1217:
1215:
1209:
1207:
1204:
1197:
1196:
1193:
1189:
1183:
1179:
1173:
1169:
1167:
1163:
1161:
1157:
1155:
1151:
1149:
1145:
1143:
1139:
1138:
1135:
1130:
1122:
1117:
1115:
1110:
1108:
1103:
1102:
1099:
1088:
1084:
1080:
1076:
1072:
1068:
1064:
1060:
1056:
1049:
1047:
1045:
1041:
1036:
1032:
1028:
1024:
1020:
1016:
1012:
1008:
1004:
1000:
993:
990:
985:
981:
977:
973:
969:
965:
961:
957:
953:
949:
945:
938:
935:
930:
926:
922:
918:
913:
908:
904:
900:
896:
892:
888:
880:
877:
873:
872:
867:
863:
859:
854:
852:
848:
843:
839:
834:
829:
825:
821:
817:
813:
812:Palaeontology
809:
802:
799:
792:
784:
783:Permunda nana
780:
779:
777:
776:
771:
766:
762:
758:
757:
755:
754:
749:
744:
740:
736:
735:
733:
732:
727:
724:
720:
719:
714:
710:
706:
705:
703:
699:
695:
694:
689:
686:
685:
680:
677:
676:
671:
668:
667:
662:
659:
658:
653:
650:
646:
642:
641:
636:
633:
629:
625:
621:
620:
619:Necronectulus
615:
612:
608:
604:
603:
598:
595:
594:
589:
586:
582:
578:
574:
573:
568:
565:
561:
560:
555:
552:
551:
546:
543:
542:
537:
534:
530:
526:
525:
520:
517:
513:
512:
507:
504:
503:
498:
495:
491:
487:
486:
481:
480:
478:
475:
469:
468:
464:
462:
461:
457:
455:
454:
450:
448:
447:
443:
442:
441:
438:
437:
433:
429:
428:
424:
422:
421:
417:
416:
414:
413:
409:
408:
406:
405:
401:
399:
397:
392:
388:
384:
380:
379:
374:
373:
369:
367:
362:
358:
354:
350:
345:
343:
339:
335:
330:
328:
324:
320:
319:
314:
313:North America
311:and northern
310:
306:
305:
300:
296:
292:
288:
284:
280:
271:
266:
265:
261:
259:
258:
254:
253:
251:
248:Extan genera
246:
241:
240:C. G. Thomson
236:
233:
230:
229:
226:
223:
221:Superfamily:
220:
219:
216:
213:
210:
209:
206:
203:
200:
199:
196:
193:
190:
189:
186:
183:
180:
179:
176:
173:
170:
169:
166:
163:
160:
159:
156:
153:
150:
149:
144:
139:
135:
132:
131:
125:
121:
116:
113:
112:
107:
103:
98:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
41:
36:
22:
19:
3028:
2907:Helophoridae
2899:Georissidae
2894:Epimetopidae
2810:Pleocomidae
2794:Ochodaeidae
2778:Hybosoridae
2762:Glaphyridae
2754:Geotrupidae
2732:
2729:Belohinidae
2690:Eucinetidae
2681:
2678:Decliniidae
2651:
2609:Phengodidae
2548:Cantharidae
2503:Dascillidae
2495:Dascilloidea
2477:Psephenidae
2469:Lutrochidae
2461:Limnichidae
2374:Buprestidae
2366:Buprestoidea
2306:Synchroidae
2290:Scraptiidae
2282:Salpingidae
2245:Prostomidae
2232:Oedemeridae
2208:Mordellidae
2126:Lymexylidae
2118:Lymexyloidea
2068:Attelabidae
2060:Anthribidae
2018:Smicripidae
1982:Phalacridae
1966:Nitidulidae
1961:Myraboliidae
1953:Monotomidae
1944:
1925:Kateretidae
1902:Cyclaxyridae
1862:
1859:Agapythidae
1823:Latridiidae
1773:Cerylonidae
1760:Anamorphidae
1694:Phycosecidae
1685:
1668:
1665:Metaxinidae
1618:Biphyllidae
1609:
1578:Orsodacnidae
1535:Cucujiformia
1468:Dermestidae
1354:
1338:
1277:Amphizoidae
1222:
1202:
1188:Archostemata
1172:Holometabola
1170:Superorder:
1164:Infraclass:
1062:
1058:
1002:
998:
992:
951:
947:
937:
894:
890:
879:
869:
815:
811:
801:
782:
773:
764:
760:
751:
742:
738:
731:Petrodromeus
729:
716:
708:
693:Sogdodromeus
691:
682:
675:Psacodromeus
673:
664:
655:
644:
638:
617:
600:
591:
576:
570:
564:Green Series
557:
548:
539:
522:
511:Dolichorabus
509:
500:
483:
465:
458:
451:
444:
436:Trachypachus
434:
425:
418:
410:
389:boundary of
376:
370:
366:Petrodromeus
364:
357:Cicindelidae
346:
331:
316:
304:Trachypachus
302:
282:
278:
276:
269:
264:Trachypachus
262:
255:
234:
211:(unranked):
128:
127:Specimen of
109:
18:
3182:iNaturalist
3053:Wikispecies
2951:Hydraenidae
2925:Spercheidae
2912:Hydrochidae
2876:Synteliidae
2860:Histeridae
2852:Histeroidea
2802:Passalidae
2770:Glaresidae
2627:Throscidae
2587:(fireflies)
2585:Lampyridae
2572:Eucnemidae
2564:Elateridae
2524:Elateroidea
2340:Zopheridae
2224:Mycteridae
2155:Anthicidae
2026:Sphindidae
2010:Silvanidae
1920:Hobartiidae
1907:Erotylidae
1836:Mycetaeidae
1831:Murmidiidae
1815:Euxestidae
1709:Rentoniidae
1583:Oxypeltidae
1568:Disteniidae
1401:Lepiceridae
1330:Hygrobiidae
1322:Haliplidae
1306:Dytiscidae
1285:Aspidytidae
721:Hong, 1984
684:Sinodromeus
666:Prosynactus
593:Karadromeus
559:Liassodites
541:Eodromeites
502:Denudirabus
488:Bode, 1953
412:Systolosoma
349:George Ball
334:leaf litter
318:Systolosoma
257:Systolosoma
3279:Categories
2972:Silphidae
2964:Ptiliidae
2956:Leiodidae
2943:Agyrtidae
2786:Lucanidae
2665:Scirtoidea
2601:Omethidae
2580:Jurasaidae
2429:Dryopidae
2400:Byrrhidae
2392:Byrrhoidea
2084:Brentidae
1889:Cucujidae
1871:Boganiidae
1851:Cucujoidea
1724:Thymalidae
1714:Rhadalidae
1657:Melyridae
1626:Byturidae
1588:Vesperidae
1497:Ptiniidae
1347:Noteridae
1314:Gyrinidae
1290:Carabidae
1219:Jurodidae
1211:Cupedidae
1158:Subclass:
1148:Arthropoda
1129:Coleoptera
793:References
577:Fortiseode
524:Dundorabus
485:Coreoeicos
225:Caraboidea
215:Geadephaga
201:Suborder:
195:Coleoptera
175:Arthropoda
2826:Trogidae
2820:(scarabs)
2698:Scirtidae
2673:Clambidae
2266:Pythidae
2200:Meloidae
2147:Aderidae
2050:(weevils)
1915:Helotidae
1841:Teredidae
1755:Alexiidae
1639:Cleridae
1598:Cleroidea
1431:Polyphaga
1429:Suborder
1382:Myxophaga
1380:Suborder
1335:Meruidae
1254:Suborder
1236:Ommatidae
1186:Suborder
1160:Pterygota
1140:Kingdom:
1087:248869983
1079:0891-2963
1027:0031-0301
984:237486703
976:0307-6970
929:237486696
921:0307-6970
842:0031-0239
753:Apermunda
657:Platycoxa
550:Eodromeus
533:Tithonian
440:Holarctic
396:sclerites
381:from the
378:Apermunda
353:Carabidae
161:Kingdom:
155:Eukaryota
3285:Adephaga
3104:BugGuide
3080:BioLib:
3038:Wikidata
2593:Lycidae
2437:Elmidae
2335:Ulodidae
2171:Boridae
2076:Belidae
1677:Peltidae
1270:families
1256:Adephaga
1166:Neoptera
1146:Phylum:
1142:Animalia
1131:families
1035:84935456
775:Permunda
649:Daohugou
602:Karatoma
494:Toarcian
387:Triassic
372:Permunda
361:Adephaga
327:Mesozoic
285:) are a
231:Family:
205:Adephaga
171:Phylum:
165:Animalia
151:Domain:
3262:6102864
3044:Q150582
2184:Ciidae
2092:Caridae
1154:Insecta
1152:Class:
1127:Extant
1007:Bibcode
956:Bibcode
899:Bibcode
820:Bibcode
702:Carnian
640:Novunda
628:Carnian
572:Evertus
402:Species
383:Permian
342:diurnal
338:conifer
309:Eurasia
291:beetles
191:Order:
185:Insecta
181:Class:
3259:uBio:
3239:575215
3213:678314
3200:114640
3187:178887
3109:165807
3096:200447
3083:249631
1268:Extant
1085:
1077:
1033:
1025:
982:
974:
927:
919:
840:
632:Norian
585:Aptian
391:Russia
287:family
242:, 1857
3252:69184
3226:60838
3195:IRMNG
3148:11007
3122:8KTMN
1083:S2CID
1031:S2CID
980:S2CID
925:S2CID
323:Chile
299:coxae
3221:NCBI
3208:ITIS
3174:7859
3169:GBIF
3135:8627
3091:BOLD
1075:ISSN
1023:ISSN
972:ISSN
917:ISSN
864:and
838:ISSN
645:Unda
375:and
277:The
35:PreꞒ
3130:EoL
3117:CoL
3068:ADW
1067:doi
1015:doi
964:doi
907:doi
828:doi
344:).
336:of
321:in
289:of
3281::
3249::
3236::
3223::
3210::
3197::
3184::
3171::
3158::
3145::
3132::
3119::
3106::
3093::
3070::
3055::
3040::
1081:.
1073:.
1063:35
1061:.
1057:.
1043:^
1029:.
1021:.
1013:.
1003:47
1001:.
978:.
970:.
962:.
952:46
950:.
946:.
923:.
915:.
905:.
895:46
893:.
889:.
868:,
850:^
836:.
826:.
816:55
814:.
810:.
763:(=
741:(=
704:)
329:.
85:Pg
2735:)
2731:(
2684:)
2680:(
2654:)
2650:(
1947:)
1943:(
1865:)
1861:(
1688:)
1684:(
1671:)
1667:(
1612:)
1608:(
1341:)
1337:(
1225:)
1221:(
1205:)
1201:(
1120:e
1113:t
1106:v
1089:.
1069::
1037:.
1017::
1009::
986:.
966::
958::
931:.
909::
901::
844:.
830::
822::
781:†
772:†
767:)
759:†
750:†
737:†
728:†
715:†
707:†
690:†
681:†
672:†
663:†
654:†
637:†
634:)
630:-
616:†
599:†
590:†
587:)
569:†
556:†
547:†
538:†
535:)
521:†
508:†
499:†
496:)
482:†
385:-
368:,
90:N
80:K
75:J
70:T
65:P
60:C
55:D
50:S
45:O
40:Ꞓ
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
