111:
505:
133:
1629:
2980:
549:. When recognized as a separate division, the literature uses many possible names, including Arthrophyta, Calamophyta, Sphenophyta, or Equisetophyta. Other authors regarded the same group as a class, either within a division consisting of the vascular plants or, more recently, within an expanded fern group. When ranked as a class, the group has been termed the Equisetopsida or Sphenopsida.
469:
485:
320:(Polypodiopsida) of which they form a specialized lineage. However, the division between the horsetails and the other ferns is so ancient that many botanists, especially paleobotanists, still regard this group as fundamentally separate at the higher level.
433:
at their edges, and in extant horsetails cover the spores externally - like sacs hanging from an umbrella, with its handle embedded in the axis of the cone. In extinct groups, further protection was afforded to the spores by the presence of whorls of
361:
trifurcate at the nodes, with the central branch becoming the vein of a microphyll, and the other two moving left and right to merge with the new branches of their neighbours. The vascular system itself resembles that of the vascular plants'
411:
in each segment of the stem and rhizome that grow as the plant gets taller. This contrasts with most seed plants, which grow from an apical meristem - i.e. new growth comes only from growing tips (and widening of stems).
1009:
The organisms first appear in the fossil record during the late
Devonian, a time when land plants were undergoing a rapid diversification, with roots, seeds and leaves having only just evolved. (See
541:, the relationship of this group to other living and fossil plants was considered problematic. Because of their unclear relationships, the rank botanists assigned to the horsetails varied from
604:. The consensus classification produced by the Pteridophyte Phylogeny Group in 2016 also places horsetails in the subclass Equisetidae, but in the class Polypodiopsida (ferns broadly defined).
328:
The horsetails comprise photosynthesising, "segmented", hollow stems, sometimes filled with pith. At the junction ("node", see diagram) between each segment is a whorl of
612:
The following diagram shows a likely phylogenic relationship between subclass
Equisetidae and the other fern subclasses according to the Pteridophyte Phylogeny Group.
2691:
2679:
2685:
857:
2495:
3566:
3286:
3073:
2697:
2827:
1660:
3312:
3099:
2507:
2623:
1013:) However, plants had already been on the land for almost a hundred million years, with the first evidence of land plants dating to
972:
The extant horsetails represent a tiny fraction of horsetail diversity in the past. There were three orders of the
Equisetidae. The
285:. They typically grow in wet areas, with whorls of needle-like branches radiating at regular intervals from a single vertical stem.
3435:
3260:
1500:
3461:
3299:
3086:
1171:
Rutishauser, R. (1999). "Polymerous leaf whorls in vascular plants: Developmental morphology and fuzziness of organ identities".
1373:
1116:
1356:
1331:
1298:
1240:
1053:
730:
A 2018 study by
Elgorriaga et al. suggests the relationships within the Equisetidae are as shown in the following cladogram.
3466:
3304:
3091:
379:
382:
are formed in the cortex. Due to the softer nature of the phloem, these are very rarely seen in fossil instances. In the
3052:
426:
1653:
371:
1076:
Pteridophyte
Phylogeny Group I (November 2016), "A community-derived classification for extant lycophytes and ferns",
1000:, but diversified as that group disappeared into extinction, gradually dwindling in diversity to today's single genus
465:: i.e. they change their configuration in the presence of water, helping the spores move and aiding their dispersal.
3104:
430:
3381:
3168:
1703:
422:
3317:
1698:
1010:
394:
grew outwards, producing a woody stem, and allowing the plants to grow as high as 10m. All extant species of
3571:
1782:
1772:
1755:
1646:
1533:"Origin of Equisetum: Evolution of horsetails (Equisetales) within the major euphyllophyte clade Sphenopsida"
3515:
3422:
3224:
3037:
132:
3561:
3386:
3327:
3173:
3114:
529:
The horsetails and their fossil relatives have long been recognized as distinct from other seedless
1638:
1323:
586:
17:
504:
3215:
1787:
1633:
538:
340:) with a singular vascular trace, fused into a sheath at each stem node. However, the leaves of
110:
3533:
3479:
3340:
3078:
3005:
1573:
1374:"Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants"
1372:
Pryer, K.M.; Schneider, H.; Smith, A.R.; Cran, R.; Wolf, P.G.; Hunt, J.S.; Sipes, S.D. (2001).
572:
1115:
Smith, A.R.; Pryer, K.M.; Schuettpelz, E.; Korall, P.; Schneider, H. & Wolf, P.G. (2006).
3528:
3474:
3363:
3335:
3127:
1750:
3150:
2988:
1315:
3291:
2983:
2729:
1588:
1388:
689:
367:
370:. Very rapid internode elongation results in the formation of a pith cavity and a ring of
8:
3556:
2148:
2131:
2041:
1889:
1316:
774:
454:
213:
119:
1592:
1392:
408:
2424:
2175:
2158:
2000:
1884:
1879:
1767:
1745:
1733:
1708:
1612:
1468:
1441:
1412:
1204:
1188:
1139:
1095:
247:
127:
3122:
2890:
2791:
2779:
2602:
2316:
2136:
2067:
2057:
1987:
1777:
1740:
1728:
1718:
1604:
1554:
1514:
1473:
1420:
1404:
1352:
1327:
1294:
1236:
1196:
1147:
1049:
546:
289:
1099:
2803:
2797:
2761:
2444:
2306:
2191:
2092:
1692:
1616:
1596:
1544:
1531:
Elgorriaga, A.; Escapa, I.H.; Rothwell, G.W.; Tomescu, A.M.F.; Cúneo, N.R. (2018).
1509:
1463:
1455:
1446:
1416:
1396:
1208:
1180:
1131:
1085:
927:
391:
3520:
3427:
3229:
3042:
3506:
2895:
2878:
2785:
2311:
2294:
2077:
2036:
1904:
1894:
1874:
1498:(2009), "A phylogenetic classification of the land plants to accompany APG III",
1495:
997:
981:
973:
748:
597:
576:
542:
358:
266:
238:
232:
3413:
3028:
2815:
2755:
2629:
2596:
2554:
2467:
2395:
2390:
2338:
1808:
1723:
1670:
652:
619:
564:
530:
493:
274:
157:
3373:
3160:
1192:
316:
have also been used. They are now recognized as rather close relatives of the
3550:
3200:
2905:
2749:
2712:
2669:
2582:
2558:
2375:
2097:
2072:
1954:
1899:
1491:
985:
600:(2009) treated the horsetails as subclass Equisetidae of class Equisetopsida
593:
581:
553:
350:
68:
36:
3140:
3353:
2966:
2821:
2809:
2549:
2380:
2370:
2170:
2122:
1995:
1608:
1558:
1477:
1408:
1200:
951:
872:
825:
568:
468:
383:
224:
219:
1293:(5th ed.). New York: Harper-Collins. pp. 371–387, 478, 506–514.
1258:
The origin of the land flora, a theory based upon the facts of alternation
3448:
3273:
3209:
3060:
2773:
2767:
2562:
2543:
2501:
2451:
2258:
2201:
2142:
2087:
2020:
1970:
1940:
1922:
1914:
1459:
993:
943:
760:
674:
462:
446:
442:
207:
43:
3239:
1600:
1143:
2973:
2900:
2868:
2854:
2837:
2385:
2062:
1851:
1842:
989:
497:
400:
are herbaceous, and have lost the ability to produce secondary growth.
337:
88:
53:
3453:
3065:
1549:
1532:
1351:. Washington, D. C.: Smithsonian Institution Press. pp. 241–242.
1349:
The Origin and early diversification of land plants: A cladistic study
1090:
556:, back to 2001, demonstrated that horsetails belong firmly within the
3278:
3247:
2638:
2534:
2400:
2284:
2082:
1862:
1831:
1400:
1135:
1015:
1002:
956:
947:
585:. (This distinction is necessary because of the alternative usage of
508:
489:
484:
417:
396:
345:
93:
2999:
1668:
592:
as a synonym for all land plants (Embryophyta) with rank of class.)
425:) at the tips of some stems. These cones comprise spirally arranged
3500:
3407:
3348:
3265:
3194:
3135:
3022:
2910:
2343:
2326:
1762:
1628:
1184:
977:
931:
278:
83:
78:
63:
58:
48:
3440:
3252:
1275:. New York and London: McGraw-Hill Book Company. pp. 110–115.
2863:
2566:
2365:
1822:
404:
363:
98:
73:
1530:
575:
era, and considered the ferns (monilophytes), to comprise four
458:
387:
2257:
1322:(3rd ed.). New York: W. H. Freeman and Company. pp.
1075:
2849:
2262:
1960:
1674:
560:
435:
375:
192:
144:
2634:
2353:
1114:
557:
534:
453:
appear in the fossil record. The sporangia open by lateral
407:, from which roots and aerial axes emerge. The plants have
329:
317:
270:
177:
167:
1962:
1571:
1288:
1048:. Englewood Cliffs, NJ: Prentice Hall. pp. 303–305.
567:. Smith et al. (2006) carried out the first higher-level
1440:
Christenhusz, Maarten J.M. & Chase, Mark W. (2014).
1371:
1235:(2nd ed.). Cambridge: Cambridge University Press.
403:
The underground parts of the plants consist of jointed
1676:
1572:
Wellman, C.H.; Osterloff, P.L.; Mohiuddin, U. (2003).
1289:
Bold, H.C.; Alexopoulos, C.J.; Delevoryas, T. (1987).
922:
According to the study, the age of the crown group of
457:
to release the spores. The spores bear characteristic
354:, in which the leaves are broad with branching veins.
942:
288:
1110:
1108:
438:- big pointed microphylls protruding from the cone.
2484:
1526:
1524:
500:(spore capsules) full of spores, can be discerned.
2433:
2157:
1439:
1307:
1105:
3548:
1986:
461:, distinctive spring-like attachments which are
2665:
1521:
1230:
537:(Polypodiopsida). Before the advent of modern
2615:
2264:
1260:. New York: Hafner publishing Co. p. 381.
1806:
1654:
1435:
1433:
1313:
1043:
579:, with the horsetails as class Equisetopsida
348:, as evidenced by early fossil forms such as
1442:"Trends and concepts in fern classification"
1346:
1340:
1273:Morphology of vascular plants (Lower Groups)
1226:
1224:
1222:
1220:
1218:
1039:
1037:
1035:
1033:
988:understory, and prospered until the mid and
1565:
1490:
1318:Morphology and evolution of vascular plants
1284:
1282:
1231:Stewart, W.N. & Rothwell, G.W. (1993).
1170:
2979:
1661:
1647:
1484:
1430:
1365:
1046:The biology and evolution of fossil plants
109:
2533:
2282:
1548:
1513:
1467:
1264:
1215:
1089:
1044:Taylor, T.N. & Taylor., E.L. (1993).
1030:
1501:Botanical Journal of the Linnean Society
1279:
503:
483:
467:
1574:"Fragments of the earliest land plants"
1233:Paleobotany and the evolution of plants
1173:International Journal of Plant Sciences
277:with a fossil record going back to the
3567:Extant Late Devonian first appearances
3549:
1071:
1069:
1067:
1065:
3004:
3003:
2256:
1642:
1270:
1255:
1249:
1164:
386:, secondary xylem (but not secondary
308:. When treated as a class, the names
3328:47c22b52-d9bb-4277-ae11-7818b1b4ff4b
3115:224f5298-14bc-4a5b-ba60-37fe9effda5d
3053:87c43432-6713-40e9-a4e3-0dd5677f312e
1314:Gifford, E.M.; Foster, A.S. (1988).
1078:Journal of Systematics and Evolution
374:formed by disruption of the primary
1117:"A classification for extant ferns"
1062:
415:Horsetails bear cones (technically
344:probably arose by the reduction of
13:
946:. This order consists of a single
366:, which evolved independently and
14:
3583:
1347:Kenrick, P.; Crane, P.R. (1997).
524:
2978:
1627:
1515:10.1111/j.1095-8339.2009.01002.x
967:
571:classification published in the
515:, terminal on an unbranched stem
131:
41:
2989:Current definitions of Plantae
1291:Morphology of Plants and Fungi
1011:Evolutionary history of plants
984:were a dominant member of the
937:
930:, and most probably up to the
323:
1:
1023:
281:. They are commonly known as
607:
332:. In the only extant genus
7:
976:first appeared in the late
519:
292:of spore plants and called
10:
3588:
1537:American Journal of Botany
441:The extant horsetails are
336:, these are small leaves (
15:
3490:
3397:
3184:
3012:
2961:
2877:
2836:
2725:
2710:
2661:
2652:
2611:
2591:
2580:
2529:
2520:
2480:
2465:
2461:
2421:
2417:
2352:
2325:
2293:
2278:
2274:
2252:
2200:
2184:
2166:
2120:
2116:
2050:
2029:
2017:
2013:
1982:
1953:
1931:
1913:
1861:
1841:
1821:
1817:
1802:
1686:
870:
854:
847:
822:
801:
794:
787:
771:
764:
745:
738:
687:
672:
665:
650:
643:
632:Equisetidae (horsetails)
630:
623:
253:
246:
204:
199:
128:Scientific classification
126:
117:
108:
29:
488:Cross-section through a
18:Equisetopsida sensu lato
1018: million years ago
996:existed alongside the
964:has about 20 species.
926:dates at least to the
573:molecular phylogenetic
516:
501:
481:
451:Calamostachys casheana
390:) was secreted as the
378:. Similar spaces, the
3529:Paleobiology Database
3475:Paleobiology Database
3336:Paleobiology Database
1256:Bower, F.O. (1959) .
554:phylogenetic analysis
507:
487:
480:L = fused microphylls
471:
409:intercalary meristems
3572:Late Devonian plants
3050:Euro+Med PlantBase:
2984:List of plant orders
2864:Pinopsida (conifers)
2850:Cycadopsida (cycads)
2842:(living gymnosperms)
2339:Leiosporocerotopsida
2058:Nephroselmidophyceae
1271:Eames, A.J. (1936).
478:B = branch in whorl,
273:(ferns), a group of
2692:Archaeopteridopsida
2425:Polysporangiophytes
2149:Klebsormidiophyceae
2132:Mesostigmatophyceae
2078:Chlorodendrophyceae
2042:Pyramimonadophyceae
1996:Prasinodermophyceae
1890:Compsopogonophyceae
1601:10.1038/nature01884
1593:2003Natur.425..282W
1393:2001Natur.409..618S
775:Archaeocalamitaceae
265:is one of the four
214:Archaeocalamitaceae
120:Equisetum telmateia
2883:(flowering plants)
2804:Lyginopteridopsida
2597:Eophyllophytopsida
2550:Zosterophyllopsida
2488:Paratracheophytes*
2266:sensu strictissimo
2176:Coleochaetophyceae
2159:Phragmoplastophyta
2001:Palmophyllophyceae
1923:Glaucocystophyceae
1885:Stylonematophyceae
1880:Porphyridiophyceae
1669:Classification of
1460:10.1093/aob/mct299
517:
502:
482:
3544:
3543:
3123:Open Tree of Life
3006:Taxon identifiers
2997:
2996:
2957:
2956:
2953:
2952:
2949:
2948:
2945:
2944:
2941:
2940:
2937:
2936:
2933:
2932:
2929:
2928:
2925:
2924:
2921:
2920:
2891:Basal angiosperms
2792:Gigantopteridales
2741:seed plant groups
2739:and other extinct
2706:
2705:
2680:Noeggerathiopsida
2648:
2647:
2603:Trimerophytopsida
2576:
2575:
2516:
2515:
2472:(vascular plants)
2437:Protracheophytes*
2413:
2412:
2409:
2408:
2376:Andreaeobryopsida
2317:Jungermanniopsida
2248:
2247:
2244:
2243:
2240:
2239:
2236:
2235:
2232:
2231:
2228:
2227:
2224:
2223:
2220:
2219:
2137:Chlorokybophyceae
2112:
2111:
2108:
2107:
2068:Chloropicophyceae
2009:
2008:
1988:Prasinodermophyta
1949:
1948:
1587:(6955): 282–285.
1550:10.1002/ajb2.1125
1387:(6820): 618–621.
1358:978-1-56098-730-7
1333:978-0-7167-1946-5
1300:978-0-06-040839-8
1242:978-0-521-38294-6
1091:10.1111/jse.12229
1055:978-0-13-651589-0
954:, with one genus
919:
918:
910:
909:
901:
900:
892:
891:
883:
882:
836:
835:
727:
726:
718:
717:
709:
708:
700:
699:
539:molecular studies
380:vallecular canals
260:
259:
195:
22:Subclass of ferns
3579:
3562:Plant subclasses
3537:
3536:
3524:
3523:
3511:
3510:
3509:
3483:
3482:
3470:
3469:
3457:
3456:
3444:
3443:
3431:
3430:
3418:
3417:
3416:
3390:
3389:
3377:
3376:
3367:
3366:
3357:
3356:
3344:
3343:
3331:
3330:
3321:
3320:
3308:
3307:
3295:
3294:
3282:
3281:
3269:
3268:
3256:
3255:
3243:
3242:
3233:
3232:
3220:
3219:
3218:
3205:
3204:
3203:
3177:
3176:
3164:
3163:
3154:
3153:
3144:
3143:
3131:
3130:
3118:
3117:
3108:
3107:
3095:
3094:
3082:
3081:
3069:
3068:
3056:
3055:
3046:
3045:
3033:
3032:
3031:
3001:
3000:
2982:
2981:
2896:Core angiosperms
2884:
2843:
2838:Acrogymnospermae
2816:Peltaspermopsida
2798:Glossopteridales
2780:Corystospermales
2762:Callistophytales
2742:
2734:
2728:
2723:
2722:
2719:
2718:
2686:Aneurophytopsida
2668:
2663:
2662:
2659:
2658:
2655:
2613:
2612:
2589:
2588:
2585:
2544:Barinophytopsida
2531:
2530:
2527:
2526:
2523:
2487:
2482:
2481:
2478:
2477:
2474:
2473:
2463:
2462:
2445:Horneophytopsida
2436:
2431:
2430:
2427:
2419:
2418:
2359:
2344:Anthocerotopsida
2332:
2327:Anthocerotophyta
2307:Haplomitriopsida
2300:
2291:
2290:
2287:
2280:
2279:
2276:
2275:
2268:
2254:
2253:
2207:
2198:
2197:
2192:Zygnematophyceae
2182:
2181:
2164:
2163:
2155:
2154:
2125:
2118:
2117:
2093:Trebouxiophyceae
2063:Picocystophyceae
2027:
2026:
2023:
2015:
2014:
1984:
1983:
1980:
1979:
1976:
1966:
1868:
1819:
1818:
1815:
1814:
1811:
1804:
1803:
1715:
1695:
1680:
1663:
1656:
1649:
1640:
1639:
1632:Data related to
1631:
1621:
1620:
1578:
1569:
1563:
1562:
1552:
1543:(8): 1286–1303.
1528:
1519:
1518:
1517:
1496:Reveal, James L.
1488:
1482:
1481:
1471:
1447:Annals of Botany
1437:
1428:
1427:
1425:
1419:. Archived from
1401:10.1038/35054555
1378:
1369:
1363:
1362:
1344:
1338:
1337:
1321:
1311:
1305:
1304:
1286:
1277:
1276:
1268:
1262:
1261:
1253:
1247:
1246:
1228:
1213:
1212:
1168:
1162:
1161:
1159:
1158:
1152:
1146:. Archived from
1136:10.2307/25065646
1121:
1112:
1103:
1102:
1093:
1073:
1060:
1059:
1041:
1019:
928:Early Cretaceous
850:
849:
797:
796:
790:
789:
767:
766:
741:
740:
734:
733:
668:
667:
646:
645:
626:
625:
616:
615:
496:, with attached
472:Vegetative stem:
449:species such as
359:vascular bundles
191:
136:
135:
113:
103:
40:
33:Temporal range:
27:
26:
3587:
3586:
3582:
3581:
3580:
3578:
3577:
3576:
3547:
3546:
3545:
3540:
3532:
3527:
3519:
3514:
3505:
3504:
3499:
3486:
3478:
3473:
3465:
3460:
3452:
3447:
3439:
3434:
3426:
3421:
3412:
3411:
3406:
3393:
3385:
3380:
3372:
3370:
3362:
3360:
3352:
3347:
3339:
3334:
3326:
3324:
3316:
3311:
3303:
3298:
3290:
3285:
3277:
3272:
3264:
3259:
3251:
3246:
3238:
3236:
3228:
3223:
3214:
3213:
3208:
3199:
3198:
3193:
3180:
3172:
3167:
3159:
3157:
3149:
3147:
3139:
3134:
3126:
3121:
3113:
3111:
3103:
3098:
3090:
3085:
3077:
3072:
3064:
3059:
3051:
3049:
3041:
3036:
3027:
3026:
3021:
3008:
2998:
2993:
2917:
2882:
2881:
2873:
2841:
2840:
2832:
2786:Czekanowskiales
2740:
2738:
2736:
2735:
2732:
2726:
2716:
2715:
2711:
2702:
2666:
2653:
2644:
2616:Moniliformopses
2607:
2581:
2572:
2522:Eutracheophytes
2521:
2512:
2485:
2471:
2470:
2466:
2457:
2434:
2422:
2405:
2396:Polytrichopsida
2391:Tetraphidopsida
2357:
2356:
2348:
2330:
2329:
2321:
2312:Marchantiopsida
2298:
2297:
2295:Marchantiophyta
2283:
2270:
2216:
2205:
2204:
2196:
2180:
2153:
2121:
2104:
2046:
2037:Mamiellophyceae
2018:
2005:
1974:
1968:
1958:
1957:
1945:
1927:
1909:
1905:Florideophyceae
1895:Rhodellophyceae
1875:Cyanidiophyceae
1866:
1865:
1857:
1837:
1807:
1798:
1797:
1713:
1691:
1682:
1667:
1636:at Wikispecies
1625:
1624:
1576:
1570:
1566:
1529:
1522:
1489:
1485:
1438:
1431:
1423:
1376:
1370:
1366:
1359:
1345:
1341:
1334:
1312:
1308:
1301:
1287:
1280:
1269:
1265:
1254:
1250:
1243:
1229:
1216:
1169:
1165:
1156:
1154:
1150:
1119:
1113:
1106:
1074:
1063:
1056:
1042:
1031:
1026:
1014:
998:Sphenophyllales
982:Sphenophyllales
974:Pseudoborniales
970:
940:
920:
911:
902:
893:
884:
858:Neocalamitaceae
837:
749:Sphenophyllales
728:
719:
710:
701:
610:
565:vascular plants
531:vascular plants
527:
522:
494:sporangiophores
479:
477:
475:
473:
427:sporangiophores
326:
275:vascular plants
239:Sphenophyllales
233:Pseudoborniales
190:
130:
104:
102:
101:
96:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
35:
34:
31:
23:
20:
12:
11:
5:
3585:
3575:
3574:
3569:
3564:
3559:
3542:
3541:
3539:
3538:
3525:
3512:
3496:
3494:
3488:
3487:
3485:
3484:
3471:
3458:
3445:
3432:
3419:
3403:
3401:
3395:
3394:
3392:
3391:
3378:
3368:
3358:
3345:
3332:
3322:
3309:
3296:
3283:
3270:
3257:
3244:
3234:
3221:
3206:
3190:
3188:
3182:
3181:
3179:
3178:
3165:
3155:
3145:
3132:
3119:
3109:
3096:
3083:
3070:
3057:
3047:
3034:
3018:
3016:
3010:
3009:
2995:
2994:
2992:
2991:
2986:
2976:
2970:
2962:
2959:
2958:
2955:
2954:
2951:
2950:
2947:
2946:
2943:
2942:
2939:
2938:
2935:
2934:
2931:
2930:
2927:
2926:
2923:
2922:
2919:
2918:
2916:
2915:
2914:
2913:
2908:
2903:
2893:
2887:
2885:
2875:
2874:
2872:
2871:
2866:
2861:
2852:
2846:
2844:
2834:
2833:
2831:
2830:
2824:
2822:Pentoxylopsida
2818:
2812:
2806:
2800:
2794:
2788:
2782:
2776:
2770:
2764:
2758:
2756:Calamopityales
2752:
2745:
2743:
2720:
2713:Spermatophytes
2708:
2707:
2704:
2703:
2701:
2700:
2694:
2688:
2682:
2675:
2673:
2670:Progymnosperms
2656:
2650:
2649:
2646:
2645:
2643:
2642:
2635:Polypodiopsida
2632:
2630:Cladoxylopsida
2626:
2619:
2617:
2609:
2608:
2606:
2605:
2599:
2592:
2586:
2583:Euphyllophytes
2578:
2577:
2574:
2573:
2571:
2570:
2555:Lycopodiopsida
2552:
2546:
2539:
2537:
2524:
2518:
2517:
2514:
2513:
2511:
2510:
2504:
2498:
2496:Cooksoniopsida
2491:
2489:
2475:
2459:
2458:
2456:
2455:
2447:
2440:
2438:
2428:
2415:
2414:
2411:
2410:
2407:
2406:
2404:
2403:
2398:
2393:
2388:
2386:Oedipodiopsida
2383:
2378:
2373:
2368:
2362:
2360:
2350:
2349:
2347:
2346:
2341:
2335:
2333:
2323:
2322:
2320:
2319:
2314:
2309:
2303:
2301:
2288:
2272:
2271:
2250:
2249:
2246:
2245:
2242:
2241:
2238:
2237:
2234:
2233:
2230:
2229:
2226:
2225:
2222:
2221:
2218:
2217:
2215:
2214:
2210:
2208:
2195:
2194:
2188:
2186:
2179:
2178:
2173:
2167:
2161:
2152:
2151:
2146:
2139:
2134:
2128:
2126:
2114:
2113:
2110:
2109:
2106:
2105:
2103:
2102:
2101:
2100:
2095:
2090:
2080:
2075:
2070:
2065:
2060:
2054:
2052:
2048:
2047:
2045:
2044:
2039:
2033:
2031:
2030:Prasinophytina
2024:
2011:
2010:
2007:
2006:
2004:
2003:
1998:
1992:
1990:
1977:
1951:
1950:
1947:
1946:
1944:
1943:
1937:
1935:
1933:incertae sedis
1929:
1928:
1926:
1925:
1919:
1917:
1911:
1910:
1908:
1907:
1902:
1897:
1892:
1887:
1882:
1877:
1871:
1869:
1859:
1858:
1856:
1855:
1847:
1845:
1839:
1838:
1836:
1835:
1827:
1825:
1812:
1809:Archaeplastida
1800:
1799:
1796:
1795:
1794:
1793:
1792:
1791:
1788:Mesomycetozoea
1785:
1780:
1770:
1760:
1759:
1758:
1753:
1748:
1743:
1738:
1737:
1736:
1724:Diaphoretickes
1721:
1716:
1711:
1706:
1701:
1696:
1688:
1687:
1684:
1683:
1671:Archaeplastida
1666:
1665:
1658:
1651:
1643:
1623:
1622:
1564:
1520:
1508:(2): 122–127,
1492:Chase, Mark W.
1483:
1454:(4): 571–594.
1429:
1426:on 2009-09-02.
1364:
1357:
1339:
1332:
1306:
1299:
1278:
1263:
1248:
1241:
1214:
1193:10.1086/314221
1185:10.1086/314221
1163:
1130:(3): 705–731.
1104:
1084:(6): 563–603,
1061:
1054:
1028:
1027:
1025:
1022:
969:
966:
939:
936:
917:
916:
913:
912:
908:
907:
904:
903:
899:
898:
895:
894:
890:
889:
886:
885:
881:
880:
877:
876:
869:
866:
865:
862:
861:
853:
848:
846:
843:
842:
839:
838:
834:
833:
830:
829:
821:
818:
817:
814:
813:
805:Paracalamitina
800:
795:
793:
788:
786:
783:
782:
779:
778:
770:
765:
763:
757:
756:
753:
752:
744:
739:
737:
732:
725:
724:
721:
720:
716:
715:
712:
711:
707:
706:
703:
702:
698:
697:
694:
693:
686:
683:
682:
679:
678:
671:
666:
664:
661:
660:
657:
656:
653:Ophioglossidae
649:
644:
642:
639:
638:
635:
634:
629:
624:
622:
620:Polypodiopsida
614:
609:
606:
587:Equisetopsida
533:, such as the
526:
525:Classification
523:
521:
518:
476:I = internode,
445:, but extinct
372:carinal canals
325:
322:
271:Polypodiopsida
258:
257:
251:
250:
244:
243:
242:
241:
235:
229:
228:
227:
222:
216:
202:
201:
197:
196:
185:
181:
180:
178:Polypodiopsida
175:
171:
170:
168:Polypodiophyta
165:
161:
160:
155:
148:
147:
142:
138:
137:
124:
123:
115:
114:
106:
105:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
42:
32:
21:
9:
6:
4:
3:
2:
3584:
3573:
3570:
3568:
3565:
3563:
3560:
3558:
3555:
3554:
3552:
3535:
3530:
3526:
3522:
3517:
3513:
3508:
3502:
3498:
3497:
3495:
3493:
3489:
3481:
3476:
3472:
3468:
3463:
3459:
3455:
3450:
3446:
3442:
3437:
3433:
3429:
3424:
3420:
3415:
3409:
3405:
3404:
3402:
3400:
3399:Equisetophyta
3396:
3388:
3383:
3379:
3375:
3369:
3365:
3359:
3355:
3350:
3346:
3342:
3337:
3333:
3329:
3323:
3319:
3314:
3310:
3306:
3301:
3297:
3293:
3288:
3284:
3280:
3275:
3271:
3267:
3262:
3258:
3254:
3249:
3245:
3241:
3235:
3231:
3226:
3222:
3217:
3216:Equisetopsida
3211:
3207:
3202:
3196:
3192:
3191:
3189:
3187:
3186:Equisetopsida
3183:
3175:
3170:
3166:
3162:
3156:
3152:
3146:
3142:
3137:
3133:
3129:
3124:
3120:
3116:
3110:
3106:
3101:
3097:
3093:
3088:
3084:
3080:
3075:
3071:
3067:
3062:
3058:
3054:
3048:
3044:
3039:
3035:
3030:
3024:
3020:
3019:
3017:
3015:
3011:
3007:
3002:
2990:
2987:
2985:
2977:
2975:
2971:
2968:
2964:
2963:
2960:
2912:
2909:
2907:
2904:
2902:
2899:
2898:
2897:
2894:
2892:
2889:
2888:
2886:
2880:
2876:
2870:
2867:
2865:
2862:
2860:
2858:
2853:
2851:
2848:
2847:
2845:
2839:
2835:
2829:
2825:
2823:
2819:
2817:
2813:
2811:
2807:
2805:
2801:
2799:
2795:
2793:
2789:
2787:
2783:
2781:
2777:
2775:
2771:
2769:
2765:
2763:
2759:
2757:
2753:
2751:
2750:Bennettitales
2747:
2746:
2744:
2731:
2730:Pteridosperms
2724:
2721:
2717:(seed plants)
2714:
2709:
2699:
2698:Protopityales
2695:
2693:
2689:
2687:
2683:
2681:
2677:
2676:
2674:
2671:
2664:
2660:
2657:
2651:
2640:
2636:
2633:
2631:
2627:
2625:
2621:
2620:
2618:
2614:
2610:
2604:
2600:
2598:
2594:
2593:
2590:
2587:
2584:
2579:
2568:
2564:
2560:
2556:
2553:
2551:
2547:
2545:
2541:
2540:
2538:
2536:
2532:
2528:
2525:
2519:
2509:
2505:
2503:
2499:
2497:
2493:
2492:
2490:
2483:
2479:
2476:
2469:
2468:Tracheophytes
2464:
2460:
2454:
2453:
2448:
2446:
2442:
2441:
2439:
2432:
2429:
2426:
2420:
2416:
2402:
2399:
2397:
2394:
2392:
2389:
2387:
2384:
2382:
2381:Andreaeopsida
2379:
2377:
2374:
2372:
2369:
2367:
2364:
2363:
2361:
2355:
2351:
2345:
2342:
2340:
2337:
2336:
2334:
2328:
2324:
2318:
2315:
2313:
2310:
2308:
2305:
2304:
2302:
2296:
2292:
2289:
2286:
2281:
2277:
2273:
2269:
2267:
2260:
2255:
2251:
2212:
2211:
2209:
2206:(land plants)
2203:
2199:
2193:
2190:
2189:
2187:
2183:
2177:
2174:
2172:
2169:
2168:
2165:
2162:
2160:
2156:
2150:
2147:
2145:
2144:
2140:
2138:
2135:
2133:
2130:
2129:
2127:
2124:
2119:
2115:
2099:
2098:Chlorophyceae
2096:
2094:
2091:
2089:
2086:
2085:
2084:
2081:
2079:
2076:
2074:
2073:Pedinophyceae
2071:
2069:
2066:
2064:
2061:
2059:
2056:
2055:
2053:
2051:Chlorophytina
2049:
2043:
2040:
2038:
2035:
2034:
2032:
2028:
2025:
2022:
2016:
2012:
2002:
1999:
1997:
1994:
1993:
1991:
1989:
1985:
1981:
1978:
1972:
1967:
1965:
1956:
1955:Viridiplantae
1952:
1942:
1939:
1938:
1936:
1934:
1930:
1924:
1921:
1920:
1918:
1916:
1912:
1906:
1903:
1901:
1900:Bangiophyceae
1898:
1896:
1893:
1891:
1888:
1886:
1883:
1881:
1878:
1876:
1873:
1872:
1870:
1864:
1860:
1854:
1853:
1849:
1848:
1846:
1844:
1840:
1834:
1833:
1829:
1828:
1826:
1824:
1820:
1816:
1813:
1810:
1805:
1801:
1789:
1786:
1784:
1781:
1779:
1776:
1775:
1774:
1771:
1769:
1766:
1765:
1764:
1761:
1757:
1754:
1752:
1751:Stramenopiles
1749:
1747:
1744:
1742:
1739:
1735:
1732:
1731:
1730:
1727:
1726:
1725:
1722:
1720:
1717:
1714:(major groups
1712:
1710:
1707:
1705:
1702:
1700:
1697:
1694:
1690:
1689:
1685:
1681:
1679:
1672:
1664:
1659:
1657:
1652:
1650:
1645:
1644:
1641:
1637:
1635:
1634:Equisetopsida
1630:
1618:
1614:
1610:
1606:
1602:
1598:
1594:
1590:
1586:
1582:
1575:
1568:
1560:
1556:
1551:
1546:
1542:
1538:
1534:
1527:
1525:
1516:
1511:
1507:
1503:
1502:
1497:
1493:
1487:
1479:
1475:
1470:
1465:
1461:
1457:
1453:
1449:
1448:
1443:
1436:
1434:
1422:
1418:
1414:
1410:
1406:
1402:
1398:
1394:
1390:
1386:
1382:
1375:
1368:
1360:
1354:
1350:
1343:
1335:
1329:
1325:
1320:
1319:
1310:
1302:
1296:
1292:
1285:
1283:
1274:
1267:
1259:
1252:
1244:
1238:
1234:
1227:
1225:
1223:
1221:
1219:
1210:
1206:
1202:
1198:
1194:
1190:
1186:
1182:
1179:(6): 81–103.
1178:
1174:
1167:
1153:on 2008-02-26
1149:
1145:
1141:
1137:
1133:
1129:
1125:
1118:
1111:
1109:
1101:
1097:
1092:
1087:
1083:
1079:
1072:
1070:
1068:
1066:
1057:
1051:
1047:
1040:
1038:
1036:
1034:
1029:
1021:
1017:
1012:
1007:
1005:
1004:
999:
995:
991:
990:early Permian
987:
986:Carboniferous
983:
979:
975:
968:Fossil record
965:
963:
959:
958:
953:
949:
945:
935:
933:
929:
925:
915:
914:
906:
905:
897:
896:
888:
887:
879:
878:
875:
874:
868:
867:
864:
863:
860:
859:
852:
851:
845:
844:
841:
840:
832:
831:
828:
827:
820:
819:
816:
815:
812:
810:
806:
803:A.G. clade (†
799:
798:
792:
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681:
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641:
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628:
627:
621:
618:
617:
613:
605:
603:
599:
595:
591:
590:
584:
583:
582:sensu stricto
578:
574:
570:
566:
562:
559:
555:
550:
548:
544:
540:
536:
532:
514:
510:
506:
499:
495:
491:
486:
470:
466:
464:
460:
456:
452:
448:
447:heterosporous
444:
439:
437:
432:
429:, which bear
428:
424:
420:
419:
413:
410:
406:
401:
399:
398:
393:
389:
385:
381:
377:
373:
369:
365:
360:
355:
353:
352:
351:Sphenophyllum
347:
343:
339:
335:
331:
321:
319:
315:
311:
310:Equisetopsida
307:
303:
299:
295:
294:Equisetophyta
291:
286:
284:
280:
276:
272:
268:
264:
256:
252:
249:
245:
240:
236:
234:
230:
226:
223:
221:
217:
215:
211:
210:
209:
206:
205:
203:
198:
194:
189:
186:
183:
182:
179:
176:
173:
172:
169:
166:
163:
162:
159:
158:Tracheophytes
156:
153:
150:
149:
146:
143:
140:
139:
134:
129:
125:
122:
121:
116:
112:
107:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
38:
37:Late Devonian
28:
25:
19:
3491:
3398:
3185:
3013:
2967:paraphyletic
2879:Angiospermae
2856:
2855:Ginkgoales (
2828:Petriellales
2810:Medullosales
2737:(seed ferns)
2450:
2371:Sphagnopsida
2366:Takakiopsida
2299:(liverworts)
2265:
2259:Embryophytes
2213:(see below↓)
2171:Charophyceae
2141:
2123:Streptophyta
1975:land plants)
1963:
1932:
1850:
1843:Rhodelphidia
1830:
1773:Opisthokonta
1677:
1626:
1584:
1580:
1567:
1540:
1536:
1505:
1499:
1486:
1451:
1445:
1421:the original
1384:
1380:
1367:
1348:
1342:
1317:
1309:
1290:
1272:
1266:
1257:
1251:
1232:
1176:
1172:
1166:
1155:. Retrieved
1148:the original
1127:
1123:
1081:
1077:
1045:
1008:
1001:
971:
961:
955:
952:Equisetaceae
941:
923:
921:
873:Equisetaceae
871:
855:
826:Calamitaceae
823:
809:Cruciaetheca
808:
804:
802:
772:
746:
729:
690:Polypodiidae
688:
673:
651:
631:
611:
601:
588:
580:
569:pteridophyte
551:
528:
512:
450:
440:
416:
414:
402:
395:
384:Calamitaceae
368:convergently
356:
349:
341:
333:
327:
313:
309:
305:
301:
297:
293:
287:
282:
262:
261:
254:
225:Equisetaceae
220:Calamitaceae
187:
151:
118:
24:
3492:Sphenophyta
3449:iNaturalist
3274:iNaturalist
3210:Wikispecies
3061:iNaturalist
3014:Equisetidae
2774:Cordaitales
2768:Caytoniales
2654:Lignophytes
2637:(ferns and
2563:spikemosses
2502:Rhyniopsida
2452:Aglaophyton
2331:(hornworts)
2202:Embryophyta
2185:Anydrophyta
2143:Spirotaenia
2088:Ulvophyceae
2021:Chlorophyta
1971:green algae
1941:Algospongia
1915:Glaucophyta
1867:(red algae)
994:Equisetales
944:Equisetales
938:Subdivision
761:Equisetales
675:Marattiidae
463:hygroscopic
443:homosporous
338:microphylls
324:Description
314:Sphenopsida
306:Sphenophyta
302:Calamophyta
298:Arthrophyta
263:Equisetidae
208:Equisetales
188:Equisetidae
30:Equisetidae
3557:Horsetails
3551:Categories
2901:Magnoliids
2869:Gnetopsida
2639:horsetails
2567:quillworts
2559:clubmosses
2535:Lycophytes
2508:Renaliales
2285:Bryophytes
1863:Rhodophyta
1852:Rhodelphis
1157:2019-10-28
1024:References
602:sensu lato
589:sensu lato
513:E. braunii
455:dehiscence
346:megaphylls
283:horsetails
267:subclasses
255:See text.
184:Subclass:
164:Division:
16:See also:
3507:Q23044259
3141:100352359
2401:Bryopsida
2354:Bryophyta
2083:UTC clade
1832:Picomonas
1768:Amoebozoa
1746:Alveolata
1734:Cryptista
1709:Eukaryota
1003:Equisetum
962:Equisetum
957:Equisetum
948:monotypic
924:Equisetum
608:Phylogeny
509:Strobilus
498:sporangia
490:strobilus
474:N = node,
431:sporangia
423:strobilus
397:Equisetum
342:Equisetum
334:Equisetum
312:s.s. and
141:Kingdom:
39:to Recent
3501:Wikidata
3414:Q3055913
3408:Wikidata
3371:VASCAN:
3354:43000109
3349:Tropicos
3237:BioLib:
3195:Wikidata
3158:VASCAN:
3136:Tropicos
3079:11907104
3029:Q5384600
3023:Wikidata
2911:Eudicots
2906:Monocots
2624:Ibykales
2358:(mosses)
2263:Plantae
1961:Plantae
1763:Amorphea
1741:Rhizaria
1729:Hacrobia
1719:Excavata
1704:Bacteria
1675:Plantae
1609:13679913
1559:30025163
1478:24532607
1409:11214320
1201:10572024
1144:25065646
1100:39980610
978:Devonian
950:family,
932:Jurassic
547:division
520:Taxonomy
421:, sing.
418:strobili
405:rhizomes
290:division
279:Devonian
248:Synonyms
3364:9480679
3201:Q134677
3128:7577693
3105:1521258
2974:extinct
1823:Picozoa
1778:Animals
1699:Archaea
1617:4383813
1589:Bibcode
1469:3936591
1417:4367248
1389:Bibcode
1209:4658142
577:classes
552:Modern
459:elaters
392:cambium
364:eustele
237:†
231:†
218:†
212:†
200:Orders
174:Class:
145:Plantae
3521:223966
3467:500001
3454:325920
3428:223963
3387:425736
3361:uBio:
3325:NZOR:
3305:500004
3230:223519
3174:450493
3151:230563
3148:uBio:
3112:NZOR:
3092:846529
3066:764793
3043:224784
2969:groups
2857:Ginkgo
2565:&
2423:
2019:
1756:Plants
1693:Domain
1615:
1607:
1581:Nature
1557:
1494:&
1476:
1466:
1415:
1407:
1381:Nature
1355:
1330:
1326:–207.
1297:
1239:
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1199:
1191:
1142:
1098:
1052:
992:. The
980:. The
598:Reveal
436:bracts
388:phloem
330:leaves
3534:56096
3480:56313
3382:WoRMS
3341:54758
3287:IRMNG
3279:47749
3253:1EQUC
3169:WoRMS
3074:IRMNG
1973:&
1783:Fungi
1613:S2CID
1577:(PDF)
1424:(PDF)
1413:S2CID
1377:(PDF)
1205:S2CID
1189:JSTOR
1151:(PDF)
1140:JSTOR
1124:Taxon
1120:(PDF)
1096:S2CID
594:Chase
561:clade
543:order
535:ferns
376:xylem
318:ferns
193:Warm.
152:Clade
3516:APNI
3462:ITIS
3436:GBIF
3423:APNI
3318:3254
3313:NCBI
3300:ITIS
3292:1295
3261:GBIF
3248:EPPO
3240:3063
3225:APNI
3100:NCBI
3087:ITIS
3038:APNI
2972:† =
1964:s.s.
1678:s.l.
1605:PMID
1555:PMID
1474:PMID
1405:PMID
1353:ISBN
1328:ISBN
1295:ISBN
1237:ISBN
1197:PMID
1050:ISBN
596:and
558:fern
357:The
44:PreꞒ
3266:246
2261:or
1959:or
1673:or
1597:doi
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1452:113
1397:doi
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