161:
818:
721:
133:
4557:
882:(mosses and liverworts) is estimated to have occurred 479–450 million years ago, and the last common ancestor of present-day hornworts lived in middle Permian about 275 million years ago. Chromosome-scale genome sequencing of three hornwort species corroborates that stomata evolved only once during land plant evolution. It also shows that the three groups of
920:
2633:
Zhang, Jian; Fu, Xin-Xing; Li, Rui-Qi; Zhao, Xiang; Liu, Yang; Li, Ming-He; Zwaenepoel, Arthur; Ma, Hong; Goffinet, Bernard; Guan, Yan-Long; Xue, Jia-Yu; Liao, Yi-Ying; Wang, Qing-Feng; Wang, Qing-Hua; Wang, Jie-Yu; Zhang, Guo-Qiang; Wang, Zhi-Wen; Jia, Yu; Wang, Mei-Zhi; Dong, Shan-Shan; Yang,
789:(closest to the interior of the gametophyte), is a foot. This is a globular group of cells that receives nutrients from the parent gametophyte, on which the sporophyte will spend its entire existence. In the middle of the sporophyte (just above the foot), is a
2286:
Qiu, Yin-Long; Li, Libo; Wang, Bin; Chen, Zhiduan; Knoop, Volker; Groth-Malonek, Milena; Dombrovska, Olena; Lee, Jungho; Kent, Livija; Rest, Joshua; Estabrook, George F.; Hendry, Tory A.; Taylor, David W.; Testa, Christopher M.; Ambros, Mathew (2006-10-17).
476:, which is both a food storing organ and enables a more efficient photosynthesis, has evolved independently five to six times in hornworts and is present in half of the roughly 200 species. It is formed by the fusion of the chloroplast with other
603:
also differ from other hornworts in having a reduced sporophyte only a few millimeters tall. The sporophyte in hornworts is unique among bryophytes in being long-lived with a persistent photosynthetic capacity. The sporophyte lacks an
671:
spore. The spores can be yellow, brown or green. Yellow and brown spores have a thicker wall and contain oils that both protect against desiccation and function as a nutrient storage, allowing them to survive for years. The species
688:
have short-lived spores with thin and colorless walls that appear green due to the presence of a chloroplast. In most species, there is a single cell inside the spore, and a slender extension of this cell called the
401:
Hornworts may be found worldwide, though they tend to grow only in places that are damp or humid. Some species grow in large numbers as tiny weeds in the soil of gardens and cultivated fields. Large tropical and
898:
inducible B gene (LCIB), which is also found in some species of algae. Because the diffusion rate of carbon dioxide is 10,000-fold higher in air than in water, aquatic algae require a mechanism to concentrate
2346:
Cooke, Todd J; Poli, DorothyBelle; Cohen, Jerry D (2003). "Did auxin play a crucial role in the evolution of novel body plans during the Late
Silurian-Early Devonian radiation of land plants?".
2912:
D. Christine
Cargill; Karen S. Renzaglia; Juan Carlos Villarreal; R. Joel Duff (2005), "Generic concepts within hornworts: Historical review, contemporary insights and future directions",
2757:
Frangedakis, Eftychios; Shimamura, Masaki; Villarreal, Juan Carlos; Li, Fay-Wei; Tomaselli, Marta; Waller, Manuel; Sakakibara, Keiko; Renzaglia, Karen S.; Szövényi, Péter (January 2021).
978:
Among land plants, hornworts are one of the earliest-diverging lineages of the early land plant ancestors; cladistic analysis implies that the group originated prior to the
2941:
Duff, R. Joel; Juan Carlos
Villarreal; D. Christine Cargill; Karen S. Renzaglia (2007). "Progress and challenges toward a phylogeny and classification of the hornworts".
950:). However, the most recent phylogenetic evidence leans strongly towards bryophyte monophyly, and it has been proposed that hornworts are de-ranked to the original class
5622:
468:, which have more than one chloroplast per cell (polyplastidy). In the polyplastidic species, and also some of the monoplastidic species, a cellular structure called a
4892:
2634:
Jian-Fen; Jiao, Yuan-Nian; Guo, Ya-Long; Kong, Hong-Zhi; Lu, An-Ming; Yang, Huan-Ming; Zhang, Shou-Zhou; Van De Peer, Yves; Liu, Zhong-Jian; Chen, Zhi-Duan (2020).
3237:
3037:"Phylogenetic and morphological infrageneric classification of the genus Dendroceros (Dendrocerotaceae; Anthocerotophyta), with the addition of two new subgenera"
993:
Structural features that have been used in the classification of hornworts include: the anatomy of chloroplasts and their numbers within cells, the presence of a
5520:
2143:
767:(singular antheridium). Both kinds of organs develop just below the surface of the plant and are only later exposed by disintegration of the overlying cells.
3142:
Renzaglia, Karen S. & Vaughn, Kevin C. (2000). Anatomy, development, and classification of hornworts. In A. Jonathan Shaw & Bernard
Goffinet (Eds.),
415:
The total number of species is still uncertain. While there are more than 300 published species names, the actual number could be as low as 100-150 species.
520:, to invade and colonize these cavities. Such colonies of bacteria growing inside the thallus give the hornwort a distinctive blue-green color. Symbiotic
2576:
Harris, Brogan J.; Clark, James W.; Schrempf, Dominik; Szöllősi, Gergely J.; Donoghue, Philip C. J.; Hetherington, Alistair M.; Williams, Tom A. (2022).
507:-filled cavities or canals when groups of cells break down. These cavities secrete hormogonium-inducing factors (HIF) that stimulate nearby, free-living
1870:
394:-dominant life cycle, in which cells of the plant carry only a single set of genetic information; the flattened, green plant body of a hornwort is the
643:
thickenings that change shape in response to drying out; they twist and thereby help to disperse the spores. Hornwort spores are relatively large for
5481:
5507:
4598:
2050:
Davies, Kevin M.; Jibran, Rubina; Zhou, Yanfei; Albert, Nick W.; Brummell, David A.; Jordan, Brian R.; Bowman, John L.; Schwinn, Kathy E. (2020).
778:, the cell from which the sporophyte stage of the life cycle will develop. Unlike all other bryophytes, the first cell division of the zygote is
866:
with central columella not attached at the roof. However, the same form of columella is also characteristic of basal moss groups, such as the
2367:
2153:
5617:
3230:
2492:
Qiu, Y.L.; Li, L.; Wang, B.; Chen, Z.; Knoop, V.; Groth-malonek, M.; Dombrovska, O.; Lee, J.; Kent, L.; Rest, J.; et al. (2006).
3206:
3155:
2560:
2476:
484:, the key enzyme in carbon fixation. By using inorganic carbon transporters and carbonic anhydrases, up to a 50-fold increase in CO
1982:"Hornwort pyrenoids, carbon-concentrating structures, evolved and were lost at least five times during the last 100 million years"
1884:"Hornwort pyrenoids, carbon-concentrating structures, evolved and were lost at least five times during the last 100 million years"
797:. Both the central and surface cells of the capsule are sterile, but between them is a layer of cells that will divide to produce
5579:
3280:
3275:
3176:
2856:
774:
must swim from the antheridia, or else be splashed to the archegonia. When this happens, the sperm and egg cell fuse to form a
1810:
MacLeod, Alexander I.; Raval, Parth K.; Stockhorst, Simon; Knopp, Michael R.; Frangedakis, Eftychios; Gould, Sven B. (2022).
744:
between one and five centimeters in diameter, and several layers of cells in thickness. It is green or yellow-green from the
5512:
160:
3359:
3349:
3332:
3223:
2978:"Bryophyte phylogeny poster: systematics and Characteristics of Nonvascular Land Plants (Mosses, Liverworts, Hornworts)"
2398:
1756:
Stotler, Raymond E.; Barbara J. Candall-Stotler (1977). "A checklist of the liverworts and hornworts of North
America".
779:
656:
652:
5419:
4591:
894:
are more closely related to each other than to hornworts. Unlike other land plants, the hornwort genome has the low-CO
1009:
Recent studies of molecular, ultrastructural, and morphological data have yielded a new classification of hornworts.
990:
are a current matter of investigation, and several competing classification schemes have been published since 1988.
3147:
2578:"Divergent evolutionary trajectories of bryophytes and tracheophytes from a complex common ancestor of land plants"
2389:
5525:
701:
grows as an extension of the original germ cell. The tip continues to divide new cells, which produces a thalloid
4735:
2954:
2808:"Large-Scale Phylogenomic Analyses Reveal the Monophyly of Bryophytes and Neoproterozoic Origin of Land Plants"
2171:"How was apical growth regulated in the ancestral land plant? Insights from the development of non-seed plants"
751:
When the gametophyte has grown to its adult size, it produces the sex organs of the hornwort. Most plants are
961:. More recently, a second class Leiosporocertotopsida has been segregated for the singularly unusual species
4820:
3364:
967:. All other hornworts remain in the class Anthocerotopsida. These two classes are divided further into five
717:
before the spore germinates. In either case, the protonema is a transitory stage in the life of a hornwort.
694:
5372:
4584:
3130:
Renzaglia, Karen S. (1978). "A comparative morphology and developmental anatomy of the
Anthocerotophyta".
148:
4268:
2441:
5048:
4698:
3327:
963:
651:
in diameter or more. The spores are polar, usually with a distinctive Y-shaped tri-radiate ridge on the
559:
embedded deep in the gametophyte. The growth of the hornwort sporophyte happens from a persistent basal
5410:
4256:
3035:
Peñaloza-Bojacá, Gabriel Felipe; Villarreal-Aguilar, Juan Carlos; Maciel-Silva, Adaíses Simone (2019).
2228:"Hornwort Stomata: Architecture and Fate Shared with 400-Million-Year-Old Fossil Plants without Leaves"
4262:
1124:
736:, which is the persistent and independent stage in the life cycle. This stage usually grows as a thin
5584:
5535:
5455:
4713:
4560:
4306:
3048:
2881:
2857:"Nuclear protein phylogenies support the monophyly of the three bryophyte groups (Bryophyta Schimp.)"
2711:
2647:
2589:
2505:
2300:
1993:
1895:
1823:
4072:
2038:
563:, in contrast to the sporophyte of moss (apical growth) and liverworts (intercalary growth). Unlike
5612:
4274:
3725:
3708:
3618:
3466:
3215:
1028:
4404:
793:
that will continue to divide and produce new cells for the third region. This third region is the
5607:
5043:
4725:
4001:
3752:
3735:
3577:
3461:
3456:
3344:
3322:
3310:
3285:
3064:
2958:
2894:
2124:
1773:
886:
share a common ancestor that branched off from the other landplants early in evolution, and that
624:
282:
263:
155:
4084:
5543:
5442:
4835:
4467:
4368:
4356:
4200:
4179:
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1962:
1923:
1853:
1550:
1354:
1159:
1082:
1812:"Loss of Plastid Developmental Genes Coincides with a Reversion to Monoplastidy in Hornworts"
755:, with both sex organs on the same plant, but some plants (even within the same species) are
5571:
5548:
5111:
4990:
4882:
4825:
4765:
4760:
4688:
4673:
4380:
4374:
4338:
4021:
3883:
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3056:
3015:
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2921:
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2778:
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2729:
2719:
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2655:
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2523:
2513:
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2406:
2355:
2318:
2308:
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2239:
2198:
2182:
2170:
2114:
2073:
2063:
2011:
2001:
1954:
1913:
1903:
1843:
1833:
1765:
1672:
1185:
972:
737:
706:
693:
germinates from the proximal side of the spore. The tip of the germ tube divides to form an
628:
431:
295:
235:
5284:
5189:
5151:
5106:
5038:
4795:
4755:
4740:
4633:
4472:
4455:
4362:
3888:
3871:
3654:
3613:
3481:
3471:
3451:
1792:
1442:
1316:
1101:
1059:
968:
636:
605:
564:
240:
230:
2226:
Renzaglia, K. S.; Villarreal, J. C.; Piatkowski, B. T.; Lucas, J. R.; Merced, A. (2017).
4576:
3104:
Grolle, Riclef (1983). "Nomina generica
Hepaticarum; references, types and synonymies".
3052:
2999:
Villareal, J. C.; Cargill, D. C.; Hagborg, A.; Söderström, L.; Renzaglia, K. S. (2010).
2715:
2651:
2593:
2509:
2304:
1997:
1899:
957:
Traditionally, there was a single class of hornworts, called
Anthocerotopsida, or older
5344:
5289:
5279:
5121:
4935:
4887:
4800:
4775:
4770:
4750:
4745:
4648:
4638:
4392:
4332:
4206:
4173:
4131:
4044:
3972:
3967:
3915:
3385:
3300:
3247:
3195:
2832:
2807:
2783:
2758:
2734:
2695:
2668:
2635:
2610:
2577:
2528:
2493:
2359:
2323:
2288:
2274:
2252:
2227:
2203:
2078:
2051:
2016:
1981:
1918:
1883:
1848:
1811:
1570:
1262:
1216:
1035:
714:
617:
508:
447:
218:
817:
623:
When the sporophyte is mature, it has a multicellular outer layer, a central rod-like
5601:
5068:
5053:
5015:
4940:
4876:
4862:
4810:
4805:
4719:
4708:
4703:
4683:
4678:
4653:
4482:
4326:
4289:
4246:
4159:
4135:
3952:
3674:
3649:
3531:
3476:
3068:
2977:
1941:
Meyer, Moritz T.; McCormick, Alistair J.; Griffiths, Howard (2016). "Will an algal CO
1871:
Hornworts: An
Overlooked Window into Carbon-Concentrating Mechanisms - Villarreal Lab
1721:
1132:
871:
710:
613:
521:
511:
144:
82:
32:
2962:
2898:
748:
in its cells, or bluish-green when colonies of cyanobacteria grow inside the plant.
5447:
5264:
5259:
5234:
5209:
5199:
5126:
4995:
4869:
4830:
4815:
4790:
4785:
4730:
4663:
4543:
4398:
4386:
4126:
3957:
3947:
3747:
3699:
3572:
1532:
1166:
867:
858:
771:
720:
403:
285:
245:
5424:
3060:
2700:
genomes illuminate the origin of land plants and the unique biology of hornworts"
5494:
5404:
5319:
5294:
5239:
5214:
5136:
5131:
5096:
5086:
4780:
4693:
4350:
4344:
4139:
4120:
4078:
4028:
3835:
3778:
3719:
3664:
3597:
3547:
3517:
3499:
3491:
3036:
1736:
1636:
1192:
998:
925:
849:
764:
760:
745:
733:
595:
556:
443:
427:
408:
395:
391:
185:
132:
57:
17:
3173:
The
Hepaticae and Anthocerotae of North America, East of the Hundredth Meridian
2601:
934:
Hornworts were traditionally considered a class within the division Bryophyta (
874:, and has been interpreted as a character common to all early land plants with
5352:
5314:
5244:
5224:
5219:
5204:
5174:
5146:
5101:
5091:
5063:
5058:
5000:
4975:
4668:
4643:
4550:
4477:
4445:
4431:
4414:
3962:
3639:
3428:
3419:
2724:
2659:
1958:
1828:
1595:
1473:
1456:
1371:
1330:
1208:
1116:
1108:
1074:
1066:
863:
853:
805:. These are released from the capsule when it splits lengthwise from the tip.
786:
648:
589:
577:
552:
532:
464:
458:
387:
379:
119:
102:
67:
5395:
3020:
2823:
2420:
2194:
2186:
1838:
759:, with separate male and female gametophytes. The female organs are known as
378:). The common name refers to the elongated horn-like structure, which is the
5468:
5269:
5249:
5141:
5020:
4955:
4945:
4930:
4925:
4906:
4658:
4215:
4111:
3977:
3861:
3659:
3439:
3408:
2518:
2494:"The deepest divergences in land plants inferred from phylogenomic evidence"
2313:
2289:"The deepest divergences in land plants inferred from phylogenomic evidence"
2068:
2006:
1908:
1619:
1200:
1140:
939:
935:
887:
883:
879:
794:
756:
752:
702:
644:
583:
526:
497:
477:
452:
107:
51:
5561:
2890:
2841:
2792:
2743:
2677:
2619:
2537:
2428:
2411:
2384:
2332:
2261:
2212:
2145:
Stress Biology of Cyanobacteria: Molecular Mechanisms to Cellular Responses
2087:
2025:
1966:
1927:
1857:
986:
known, but new species are still being discovered. The number and names of
5556:
5434:
5389:
5324:
5304:
5299:
5274:
5254:
5229:
5194:
5184:
5179:
5010:
4985:
4965:
4625:
4487:
3339:
2351:
1001:
within androecia, and the arrangement of jacket cells of the antheridia.
994:
982:, around the same time as the mosses and liverworts. There are about 200
979:
790:
560:
504:
473:
469:
123:
97:
92:
77:
72:
62:
2383:
Friedman, William E.; Moore, Richard C.; Purugganan, Michael D. (2004).
2275:
Classification of the Phylum Anthocerotophyta Stotl. & Crand.-Stotl.
2243:
919:
488:
levels can be achieved. This particular feature is very unusual in land
5329:
5309:
5005:
4980:
4970:
4960:
4916:
4853:
4440:
4143:
3942:
3399:
2128:
2039:
BTI researchers unlocking hornworts' secrets | EurekAlert! Science News
1777:
983:
904:
741:
698:
668:
612:-sensitive point of divergence with other land plants some time in the
545:
541:
481:
439:
435:
424:
112:
87:
2872:
2774:
2553:
The Origin and Early Diversification of Land Plants: A Cladistic Study
571:
on their sporophyte as most mosses do. The exceptions are the species
438:
between one and five centimeters in diameter. Hornworts have lost two
5499:
5169:
5161:
5030:
3001:"A synthesis of hornwort diversity: Patterns, causes and future work"
798:
775:
632:
516:
383:
5366:
2925:
2555:. Washington, D. C.: Smithsonian Institution Press. pp. 55–56.
2119:
2102:
1769:
2142:
Srivastava, Ashish Kumar; Rai, Amar Nath; Neilan, Brett A. (2013).
782:. Further divisions produce three basic regions of the sporophyte.
5486:
5473:
5078:
4950:
4565:
4426:
3839:
3537:
3251:
2052:"The Evolution of Flavonoid Biosynthesis: A Bryophyte Perspective"
987:
875:
802:
719:
640:
609:
568:
489:
442:
division-associated genes, ARC3 and FtsZ2, and have just a single
423:
Like all bryophytes, the dominant life phase of a hornwort is the
172:
907:
protein to function efficiently. LCIB is one component of this CO
27:
Division of non-vascular land plants with horn-shaped sporophytes
5116:
4608:
4211:
3930:
891:
493:
5370:
4580:
3833:
3219:
1764:(3). American Bryological and Lichenological Society: 405–428.
635:. The pseudo-elaters are multi-cellular, unlike the elaters of
812:
5460:
30:
The name hornwort also refers to aquatic plants of the genus
3083:
2976:
Cole, Theodor C. H.; Hilger, Hartmut H.; Goffinet, Bernard.
2103:"Symbiosis between nitrogen-fixing cyanobacteria and plants"
307:
3186:
Cryptogamic Botany, Volume II: Bryophytes and Pteridophytes
3117:
Hasegawa, J. (1994). "New classification of Anthocerotae".
862:
may represent a stem group to the clade, as it possesses a
370:
364:
355:
346:
340:
325:
319:
310:
3539:
2955:
10.1639/0007-2745(2007)110[214:PACTDA]2.0.CO;2
1980:
Villarreal, Juan Carlos; Renner, Susanne S. (2012-11-13).
1882:
Villarreal, Juan Carlos; Renner, Susanne S. (2012-11-13).
334:
942:, and hence the hornworts were given their own division,
301:
763:(singular archegonium) and the male organs are known as
3201:(3rd ed.). London: Hutchinson University Library.
829:
3253:
2759:"The hornworts: morphology, evolution and development"
2694:
Li, F-W.; Nishiyama, T.; Waller, M.; et, al. (2020).
361:
337:
316:
313:
304:
298:
2636:"The hornwort genome and early land plant evolution"
496:. They are also the only group of land plants where
367:
352:
343:
331:
322:
5379:
5342:
5160:
5077:
5029:
4914:
4905:
4852:
4624:
4454:
4413:
4302:
4287:
4242:
4229:
4192:
4157:
4110:
4097:
4061:
4042:
4010:
3998:
3929:
3902:
3870:
3859:
3777:
3761:
3734:
3697:
3627:
3606:
3594:
3563:
3530:
3508:
3490:
3438:
3418:
3398:
3383:
358:
328:
3194:
1945:-concentrating mechanism work in higher plants?".
903:in chloroplasts so as to allow the photosynthetic
4893:Archaeal Richmond Mine acidophilic nanoorganisms
2169:Fouracre, Jim P; Harrison, C Jill (2022-08-29).
450:(monoplastidy), with the exception of the genus
2498:Proceedings of the National Academy of Sciences
2293:Proceedings of the National Academy of Sciences
1986:Proceedings of the National Academy of Sciences
1888:Proceedings of the National Academy of Sciences
3841:
2462:
2460:
709:may begin dividing within the spore, becoming
4592:
3231:
2689:
2687:
8:
2936:
2934:
2453:NEW CLASSIFICATION OF ANTHOCEROTAE - J-Stage
938:). Later on, the bryophytes were considered
3132:Journal of the Hattori Botanical Laboratory
3119:Journal of the Hattori Botanical Laboratory
492:, unique to hornworts, but is common among
412:may be found growing on the bark of trees.
5367:
4911:
4849:
4621:
4617:
4599:
4585:
4577:
4556:
4299:
4239:
4235:
4189:
4165:
4107:
4103:
4058:
4054:
4039:
4007:
3995:
3867:
3856:
3852:
3830:
3774:
3758:
3740:
3731:
3694:
3603:
3591:
3560:
3556:
3395:
3391:
3380:
3238:
3224:
3216:
131:
40:
3019:
2882:1983/0b471d7e-ce54-4681-b791-1da305d9e53b
2880:
2831:
2782:
2733:
2723:
2667:
2609:
2527:
2517:
2410:
2322:
2312:
2251:
2202:
2118:
2077:
2067:
2015:
2005:
1917:
1907:
1847:
1837:
1827:
1724:and composition of the Anthocerotophyta.
659:surface ornamented with bumps or spines.
544:. These pores superficially resemble the
5623:Extant Late Cretaceous first appearances
918:
288:(land plants) constituting the division
1748:
878:. The divergence between hornworts and
2855:de Sousa, Filipe; et al. (2019).
2551:Kenrick, Paul; Peter R. Crane (1997).
1799:. Australian National Botanic Gardens.
627:running up the center, and a layer of
3188:. New York: McGraw-Hill Book Company.
705:. By contrast, species of the family
667:The life of a hornwort starts from a
581:and the three closely related genera
430:. This stage usually grows as a thin
7:
5536:448fb371-8eb0-4c66-aec5-36d20684e98e
3197:The Structure and Life of Bryophytes
2467:Chopra, R. N.; Kumra, P. K. (1988).
2385:"The evolution of plant development"
631:in between that produces spores and
2471:. New York: John Wiley & Sons.
1611:
1587:
1561:
1524:
1516:
1448:
1438:
1430:
1423:
1346:
1322:
1312:
1304:
1297:
1253:
1245:
1237:
1230:
852:hornworts only begins in the upper
732:From the protonema grows the adult
253:
25:
3082:Brinda, John C.; Atwood, John J.
2348:The Evolution of Plant Physiology
1791:Lepp, Heino (12 September 2012).
724:Life cycle of a typical hornwort
480:and is composed predominantly of
200:Stotler & Stotl.-Crand., 1977
4555:
2806:Su, Danyan; et al. (2021).
2360:10.1016/b978-012339552-8/50006-8
1947:Current Opinion in Plant Biology
816:
728:. Click on the image to enlarge.
503:Many hornworts develop internal
294:
159:
36:, in the family Ceratophyllaceae
3177:Field Museum of Natural History
2812:Molecular Biology and Evolution
456:and some species in the genera
4566:Current definitions of Plantae
2582:Nature Ecology & Evolution
930:growing on the bark of a tree.
647:, measuring between 30 and 80
1:
3061:10.1080/14772000.2019.1682080
1011:
599:, which do not have stomata.
3171:Schuster, Rudolf M. (1992).
3146:, pp. 1–20. Cambridge:
3041:Systematics and Biodiversity
2914:Australian Systematic Botany
2399:Botanical Society of America
3084:"The Bryophyte Nomenclator"
971:, each containing a single
848:While the fossil record of
5639:
5618:Turonian first appearances
3184:Smith, Gilbert M. (1938).
3148:Cambridge University Press
2602:10.1038/s41559-022-01885-x
2390:American Journal of Botany
2056:Frontiers in Plant Science
1816:Frontiers in Plant Science
1017:Class Leiosporocerotopsida
911:-concentrating mechanism.
536:. There may also be small
524:have not been reported in
29:
4848:
4620:
4616:
4611:phyla/divisions by domain
4538:
4238:
4188:
4168:
4106:
4057:
4038:
3994:
3855:
3851:
3829:
3743:
3693:
3590:
3559:
3394:
3379:
3263:
3175:. Vol. VI. Chicago:
3162:Schofield, W. B. (1985).
2725:10.1038/s41477-020-0618-2
2660:10.1038/s41477-019-0588-4
1959:10.1016/j.pbi.2016.04.009
1829:10.1101/2022.01.11.475830
1719:
1633:
1616:
1609:
1592:
1585:
1567:
1559:
1529:
1522:
1514:
1470:
1453:
1446:
1436:
1428:
1421:
1368:
1351:
1344:
1327:
1320:
1310:
1302:
1295:
1259:
1251:
1243:
1235:
269:
262:
210:
205:
156:Scientific classification
154:
139:
130:
43:
3164:Introduction to Bryology
3021:10.11646/phytotaxa.9.1.8
1839:10.3389/fpls.2022.863076
697:of cells, and the first
540:on the underside of the
514:, especially species of
4821:Thermodesulfobacteriota
2519:10.1073/pnas.0603335103
2314:10.1073/pnas.0603335103
2069:10.3389/fpls.2020.00007
2007:10.1073/pnas.1213498109
1909:10.1073/pnas.1213498109
695:octant (solid geometry)
500:are completely absent.
3193:Watson, E. V. (1971).
3166:. New York: Macmillan.
2824:10.1093/molbev/msab106
2412:10.3732/ajb.91.10.1726
2187:10.1093/plphys/kiac313
1048:Class Anthocerotopsida
946:(sometimes misspelled
931:
729:
567:, hornworts have true
5049:Neocallimastigomycota
3106:Acta Botanica Fennica
2469:Biology of Bryophytes
1797:Australian Bryophytes
1793:"What is a hornwort?"
1565:Phaeomegacerotoideae
1241:Leiosporocerotopsida
964:Leiosporoceros dussii
922:
856:, the lower Devonian
785:At the bottom of the
723:
674:Folioceros fuciformis
4714:Coprothermobacterota
4561:List of plant orders
4441:Pinopsida (conifers)
4427:Cycadopsida (cycads)
4419:(living gymnosperms)
3916:Leiosporocerotopsida
3635:Nephroselmidophyceae
1257:Leiosporocerotaceae
809:Evolutionary history
655:surface, and with a
398:stage of the plant.
213:Leiosporocerotopsida
4269:Archaeopteridopsida
4002:Polysporangiophytes
3726:Klebsormidiophyceae
3709:Mesostigmatophyceae
3655:Chlorodendrophyceae
3619:Pyramimonadophyceae
3573:Prasinodermophyceae
3467:Compsopogonophyceae
3053:2019SyBio..17..712P
2716:2020NatPl...6..259L
2652:2020NatPl...6..107Z
2594:2022NatEE...6.1634H
2510:2006PNAS..10315511Q
2354:. pp. 85–107.
2305:2006PNAS..10315511Q
2299:(42): 15511–15516.
2244:10.1104/pp.17.00156
1998:2012PNAS..10918873V
1992:(46): 18873–18878.
1900:2012PNAS..10918873V
1894:(46): 18873–18878.
1249:Leiosporocerotales
1029:Leiosporocerotaceae
573:Folioceros incurvus
390:, hornworts have a
206:Classes and orders
5044:Blastocladiomycota
4460:(flowering plants)
4381:Lyginopteridopsida
4174:Eophyllophytopsida
4127:Zosterophyllopsida
4065:Paratracheophytes*
3843:sensu strictissimo
3753:Coleochaetophyceae
3736:Phragmoplastophyta
3578:Palmophyllophyceae
3500:Glaucocystophyceae
3462:Stylonematophyceae
3457:Porphyridiophyceae
3246:Classification of
2101:Meeks, JC (1998).
1583:Dendrocerotoideae
1023:Leiosporocerotales
932:
828:. You can help by
730:
219:Leiosporocerotales
122:(but see text) to
5595:
5594:
5544:Open Tree of Life
5373:Taxon identifiers
5364:
5363:
5360:
5359:
5338:
5337:
4901:
4900:
4844:
4843:
4836:Verrucomicrobiota
4574:
4573:
4534:
4533:
4530:
4529:
4526:
4525:
4522:
4521:
4518:
4517:
4514:
4513:
4510:
4509:
4506:
4505:
4502:
4501:
4498:
4497:
4468:Basal angiosperms
4369:Gigantopteridales
4318:seed plant groups
4316:and other extinct
4283:
4282:
4257:Noeggerathiopsida
4225:
4224:
4180:Trimerophytopsida
4153:
4152:
4093:
4092:
4049:(vascular plants)
4014:Protracheophytes*
3990:
3989:
3986:
3985:
3953:Andreaeobryopsida
3894:Jungermanniopsida
3825:
3824:
3821:
3820:
3817:
3816:
3813:
3812:
3809:
3808:
3805:
3804:
3801:
3800:
3797:
3796:
3714:Chlorokybophyceae
3689:
3688:
3685:
3684:
3645:Chloropicophyceae
3586:
3585:
3565:Prasinodermophyta
3526:
3525:
3144:Bryophyte Biology
3086:. 7 December 2022
2873:10.1111/nph.15587
2775:10.1111/nph.16874
2588:(11): 1634–1643.
2442:Bryophyte Biology
2369:978-0-12-339552-8
2155:978-1-4665-0478-3
1728:
1727:
1714:
1713:
1705:
1704:
1696:
1695:
1687:
1686:
1678:
1677:
1666:
1665:
1657:
1656:
1648:
1647:
1551:Phymatocerotaceae
1544:
1543:
1520:Phymatocerotales
1503:
1502:
1494:
1493:
1485:
1484:
1410:
1409:
1401:
1400:
1392:
1391:
1383:
1382:
1355:Sphaerosporoceros
1293:Anthocerotopsida
1283:
1282:
1274:
1273:
1160:Phymatocerotaceae
1083:Sphaerosporoceros
997:, the numbers of
846:
845:
770:The biflagellate
548:of other plants.
276:
275:
201:
141:Phaeoceros laevis
16:(Redirected from
5630:
5588:
5587:
5575:
5574:
5565:
5564:
5552:
5551:
5539:
5538:
5529:
5528:
5516:
5515:
5503:
5502:
5490:
5489:
5477:
5476:
5464:
5463:
5451:
5450:
5438:
5437:
5428:
5427:
5415:
5414:
5413:
5411:Anthocerotophyta
5400:
5399:
5398:
5381:Anthocerotophyta
5368:
5112:Anthocerotophyta
4991:Hemimastigophora
4912:
4883:Nitrososphaerota
4850:
4826:Thermomicrobiota
4766:Kiritimatiellota
4761:Ignavibacteriota
4726:Deferribacterota
4689:Campylobacterota
4674:Bdellovibrionota
4622:
4618:
4601:
4594:
4587:
4578:
4559:
4558:
4473:Core angiosperms
4461:
4420:
4415:Acrogymnospermae
4393:Peltaspermopsida
4375:Glossopteridales
4357:Corystospermales
4339:Callistophytales
4319:
4311:
4305:
4300:
4296:
4295:
4263:Aneurophytopsida
4245:
4240:
4236:
4232:
4190:
4166:
4162:
4121:Barinophytopsida
4108:
4104:
4100:
4064:
4059:
4055:
4051:
4050:
4040:
4022:Horneophytopsida
4013:
4008:
4004:
3996:
3936:
3921:Anthocerotopsida
3909:
3904:Anthocerotophyta
3884:Haplomitriopsida
3877:
3868:
3864:
3857:
3853:
3845:
3831:
3784:
3775:
3769:Zygnematophyceae
3759:
3741:
3732:
3702:
3695:
3670:Trebouxiophyceae
3640:Picocystophyceae
3604:
3600:
3592:
3561:
3557:
3553:
3543:
3445:
3396:
3392:
3388:
3381:
3292:
3272:
3257:
3240:
3233:
3226:
3217:
3212:
3200:
3189:
3180:
3167:
3139:
3126:
3113:
3096:
3095:
3093:
3091:
3079:
3073:
3072:
3032:
3026:
3025:
3023:
3005:
2996:
2990:
2989:
2987:
2985:
2973:
2967:
2966:
2938:
2929:
2928:
2909:
2903:
2902:
2884:
2852:
2846:
2845:
2835:
2818:(8): 3332–3344.
2803:
2797:
2796:
2786:
2754:
2748:
2747:
2737:
2727:
2691:
2682:
2681:
2671:
2630:
2624:
2623:
2613:
2573:
2567:
2566:
2548:
2542:
2541:
2531:
2521:
2489:
2483:
2482:
2464:
2455:
2450:
2444:
2439:
2433:
2432:
2414:
2380:
2374:
2373:
2343:
2337:
2336:
2326:
2316:
2283:
2277:
2272:
2266:
2265:
2255:
2232:Plant Physiology
2223:
2217:
2216:
2206:
2175:Plant Physiology
2166:
2160:
2159:
2139:
2133:
2132:
2122:
2098:
2092:
2091:
2081:
2071:
2047:
2041:
2036:
2030:
2029:
2019:
2009:
1977:
1971:
1970:
1938:
1932:
1931:
1921:
1911:
1879:
1873:
1868:
1862:
1861:
1851:
1841:
1831:
1807:
1801:
1800:
1788:
1782:
1781:
1753:
1673:Dendrocerotaceae
1612:
1588:
1562:
1557:Dendrocerotales
1525:
1517:
1449:
1439:
1431:
1424:
1347:
1323:
1313:
1305:
1298:
1254:
1246:
1238:
1231:
1186:Dendrocerotaceae
1154:Phymatocerotales
1125:Paraphymatoceros
1120:(ca. 41 species)
1070:(ca. 83 species)
1012:
952:Anthocerotopsida
944:Anthocerotophyta
841:
838:
820:
813:
707:Dendrocerotaceae
551:The horn-shaped
377:
376:
373:
372:
369:
366:
363:
360:
357:
354:
349:
348:
345:
342:
339:
336:
333:
330:
327:
324:
321:
318:
315:
312:
309:
306:
303:
300:
290:Anthocerotophyta
246:Phymatocerotales
225:Anthocerotopsida
199:
196:Anthocerotophyta
164:
163:
135:
117:
54:
47:Temporal range:
41:
21:
18:Anthocerotopsida
5638:
5637:
5633:
5632:
5631:
5629:
5628:
5627:
5598:
5597:
5596:
5591:
5583:
5578:
5570:
5568:
5560:
5555:
5547:
5542:
5534:
5532:
5524:
5519:
5511:
5506:
5498:
5493:
5485:
5480:
5472:
5467:
5459:
5454:
5446:
5441:
5433:
5431:
5423:
5418:
5409:
5408:
5403:
5394:
5393:
5388:
5375:
5365:
5356:
5334:
5285:Gnathostomulida
5190:Xenacoelomorpha
5156:
5152:Flowering plant
5107:Marchantiophyta
5073:
5039:Chytridiomycota
5025:
4897:
4840:
4796:Planctomycetota
4756:Gemmatimonadota
4741:Elusimicrobiota
4634:Acidobacteriota
4612:
4605:
4575:
4570:
4494:
4459:
4458:
4450:
4418:
4417:
4409:
4363:Czekanowskiales
4317:
4315:
4313:
4312:
4309:
4303:
4293:
4292:
4288:
4279:
4243:
4230:
4221:
4193:Moniliformopses
4184:
4158:
4149:
4099:Eutracheophytes
4098:
4089:
4062:
4048:
4047:
4043:
4034:
4011:
3999:
3982:
3973:Polytrichopsida
3968:Tetraphidopsida
3934:
3933:
3925:
3907:
3906:
3898:
3889:Marchantiopsida
3875:
3874:
3872:Marchantiophyta
3860:
3847:
3793:
3782:
3781:
3773:
3757:
3730:
3698:
3681:
3623:
3614:Mamiellophyceae
3595:
3582:
3551:
3545:
3535:
3534:
3522:
3504:
3486:
3482:Florideophyceae
3472:Rhodellophyceae
3452:Cyanidiophyceae
3443:
3442:
3434:
3414:
3384:
3375:
3374:
3290:
3268:
3259:
3244:
3209:
3192:
3183:
3170:
3161:
3129:
3116:
3103:
3100:
3099:
3089:
3087:
3081:
3080:
3076:
3034:
3033:
3029:
3003:
2998:
2997:
2993:
2983:
2981:
2975:
2974:
2970:
2940:
2939:
2932:
2926:10.1071/sb04012
2911:
2910:
2906:
2861:New Phytologist
2854:
2853:
2849:
2805:
2804:
2800:
2763:New Phytologist
2756:
2755:
2751:
2693:
2692:
2685:
2632:
2631:
2627:
2575:
2574:
2570:
2563:
2550:
2549:
2545:
2504:(42): 15511–6.
2491:
2490:
2486:
2479:
2466:
2465:
2458:
2451:
2447:
2440:
2436:
2382:
2381:
2377:
2370:
2345:
2344:
2340:
2285:
2284:
2280:
2273:
2269:
2225:
2224:
2220:
2168:
2167:
2163:
2156:
2141:
2140:
2136:
2120:10.2307/1313353
2100:
2099:
2095:
2049:
2048:
2044:
2037:
2033:
1979:
1978:
1974:
1944:
1940:
1939:
1935:
1881:
1880:
1876:
1869:
1865:
1809:
1808:
1804:
1790:
1789:
1785:
1770:10.2307/3242017
1755:
1754:
1750:
1745:
1733:
1715:
1706:
1697:
1688:
1679:
1667:
1658:
1649:
1545:
1504:
1495:
1486:
1443:Notothyladaceae
1434:Notothyladales
1411:
1402:
1393:
1384:
1317:Anthocerotaceae
1308:Anthocerotales
1284:
1275:
1180:Dendrocerotales
1102:Notothyladaceae
1060:Anthocerotaceae
1007:
917:
910:
902:
897:
842:
836:
833:
826:needs expansion
811:
740:or ribbon-like
676:and the genera
665:
606:apical meristem
487:
472:is absent. The
434:or ribbon-like
421:
351:
297:
293:
281:are a group of
236:Dendrocerotales
198:
158:
126:
116:
115:
110:
105:
100:
95:
90:
85:
80:
75:
70:
65:
60:
49:
48:
45:
37:
28:
23:
22:
15:
12:
11:
5:
5636:
5634:
5626:
5625:
5620:
5615:
5610:
5600:
5599:
5593:
5592:
5590:
5589:
5576:
5566:
5553:
5540:
5530:
5517:
5504:
5491:
5478:
5465:
5452:
5439:
5429:
5416:
5401:
5385:
5383:
5377:
5376:
5371:
5362:
5361:
5358:
5357:
5350:
5348:
5345:Incertae sedis
5340:
5339:
5336:
5335:
5333:
5332:
5327:
5322:
5317:
5312:
5307:
5302:
5297:
5292:
5290:Micrognathozoa
5287:
5282:
5280:Acanthocephala
5277:
5272:
5267:
5262:
5257:
5252:
5247:
5242:
5237:
5232:
5227:
5222:
5217:
5212:
5207:
5202:
5197:
5192:
5187:
5182:
5177:
5172:
5166:
5164:
5158:
5157:
5155:
5154:
5149:
5144:
5139:
5134:
5129:
5124:
5122:Lycopodiophyta
5119:
5114:
5109:
5104:
5099:
5094:
5089:
5083:
5081:
5075:
5074:
5072:
5071:
5066:
5061:
5056:
5051:
5046:
5041:
5035:
5033:
5027:
5026:
5024:
5023:
5018:
5013:
5008:
5003:
4998:
4993:
4988:
4983:
4978:
4973:
4968:
4963:
4958:
4953:
4948:
4943:
4938:
4936:Ancyromonadida
4933:
4928:
4922:
4920:
4909:
4903:
4902:
4899:
4898:
4896:
4895:
4890:
4888:Thermoproteota
4885:
4880:
4873:
4866:
4858:
4856:
4846:
4845:
4842:
4841:
4839:
4838:
4833:
4828:
4823:
4818:
4813:
4808:
4803:
4801:Pseudomonadota
4798:
4793:
4788:
4783:
4778:
4776:Mycoplasmatota
4773:
4771:Lentisphaerota
4768:
4763:
4758:
4753:
4751:Fusobacteriota
4748:
4746:Fibrobacterota
4743:
4738:
4733:
4728:
4723:
4716:
4711:
4706:
4701:
4696:
4691:
4686:
4681:
4676:
4671:
4666:
4661:
4656:
4651:
4649:Armatimonadota
4646:
4641:
4639:Actinomycetota
4636:
4630:
4628:
4614:
4613:
4606:
4604:
4603:
4596:
4589:
4581:
4572:
4571:
4569:
4568:
4563:
4553:
4547:
4539:
4536:
4535:
4532:
4531:
4528:
4527:
4524:
4523:
4520:
4519:
4516:
4515:
4512:
4511:
4508:
4507:
4504:
4503:
4500:
4499:
4496:
4495:
4493:
4492:
4491:
4490:
4485:
4480:
4470:
4464:
4462:
4452:
4451:
4449:
4448:
4443:
4438:
4429:
4423:
4421:
4411:
4410:
4408:
4407:
4401:
4399:Pentoxylopsida
4395:
4389:
4383:
4377:
4371:
4365:
4359:
4353:
4347:
4341:
4335:
4333:Calamopityales
4329:
4322:
4320:
4297:
4290:Spermatophytes
4285:
4284:
4281:
4280:
4278:
4277:
4271:
4265:
4259:
4252:
4250:
4247:Progymnosperms
4233:
4227:
4226:
4223:
4222:
4220:
4219:
4212:Polypodiopsida
4209:
4207:Cladoxylopsida
4203:
4196:
4194:
4186:
4185:
4183:
4182:
4176:
4169:
4163:
4160:Euphyllophytes
4155:
4154:
4151:
4150:
4148:
4147:
4132:Lycopodiopsida
4129:
4123:
4116:
4114:
4101:
4095:
4094:
4091:
4090:
4088:
4087:
4081:
4075:
4073:Cooksoniopsida
4068:
4066:
4052:
4036:
4035:
4033:
4032:
4024:
4017:
4015:
4005:
3992:
3991:
3988:
3987:
3984:
3983:
3981:
3980:
3975:
3970:
3965:
3963:Oedipodiopsida
3960:
3955:
3950:
3945:
3939:
3937:
3927:
3926:
3924:
3923:
3918:
3912:
3910:
3900:
3899:
3897:
3896:
3891:
3886:
3880:
3878:
3865:
3849:
3848:
3834:
3827:
3826:
3823:
3822:
3819:
3818:
3815:
3814:
3811:
3810:
3807:
3806:
3803:
3802:
3799:
3798:
3795:
3794:
3792:
3791:
3787:
3785:
3772:
3771:
3765:
3763:
3756:
3755:
3750:
3744:
3738:
3729:
3728:
3723:
3716:
3711:
3705:
3703:
3691:
3690:
3687:
3686:
3683:
3682:
3680:
3679:
3678:
3677:
3672:
3667:
3657:
3652:
3647:
3642:
3637:
3631:
3629:
3625:
3624:
3622:
3621:
3616:
3610:
3608:
3607:Prasinophytina
3601:
3588:
3587:
3584:
3583:
3581:
3580:
3575:
3569:
3567:
3554:
3528:
3527:
3524:
3523:
3521:
3520:
3514:
3512:
3510:incertae sedis
3506:
3505:
3503:
3502:
3496:
3494:
3488:
3487:
3485:
3484:
3479:
3474:
3469:
3464:
3459:
3454:
3448:
3446:
3436:
3435:
3433:
3432:
3424:
3422:
3416:
3415:
3413:
3412:
3404:
3402:
3389:
3386:Archaeplastida
3377:
3376:
3373:
3372:
3371:
3370:
3369:
3368:
3365:Mesomycetozoea
3362:
3357:
3347:
3337:
3336:
3335:
3330:
3325:
3320:
3315:
3314:
3313:
3301:Diaphoretickes
3298:
3293:
3288:
3283:
3278:
3273:
3265:
3264:
3261:
3260:
3248:Archaeplastida
3245:
3243:
3242:
3235:
3228:
3220:
3214:
3213:
3207:
3190:
3181:
3168:
3159:
3140:
3127:
3114:
3098:
3097:
3074:
3047:(7): 712–727.
3027:
2991:
2968:
2949:(2): 214–243.
2943:The Bryologist
2930:
2904:
2867:(1): 565–575.
2847:
2798:
2769:(2): 735–754.
2749:
2710:(3): 259–272.
2683:
2646:(2): 107–118.
2625:
2568:
2561:
2543:
2484:
2477:
2456:
2445:
2434:
2405:): 1726–1741.
2375:
2368:
2338:
2278:
2267:
2238:(2): 788–797.
2218:
2181:(1): 100–112.
2161:
2154:
2134:
2113:(4): 266–276.
2093:
2042:
2031:
1972:
1942:
1933:
1874:
1863:
1802:
1783:
1758:The Bryologist
1747:
1746:
1744:
1741:
1740:
1739:
1732:
1729:
1726:
1725:
1717:
1716:
1712:
1711:
1708:
1707:
1703:
1702:
1699:
1698:
1694:
1693:
1690:
1689:
1685:
1684:
1681:
1680:
1676:
1675:
1669:
1668:
1664:
1663:
1660:
1659:
1655:
1654:
1651:
1650:
1646:
1645:
1642:
1641:
1632:
1629:
1628:
1625:
1624:
1615:
1610:
1608:
1605:
1604:
1601:
1600:
1591:
1586:
1584:
1580:
1579:
1576:
1575:
1571:Phaeomegaceros
1566:
1560:
1558:
1554:
1553:
1547:
1546:
1542:
1541:
1538:
1537:
1528:
1523:
1521:
1515:
1513:
1510:
1509:
1506:
1505:
1501:
1500:
1497:
1496:
1492:
1491:
1488:
1487:
1483:
1482:
1479:
1478:
1469:
1466:
1465:
1462:
1461:
1452:
1447:
1445:
1437:
1435:
1429:
1427:
1422:
1420:
1417:
1416:
1413:
1412:
1408:
1407:
1404:
1403:
1399:
1398:
1395:
1394:
1390:
1389:
1386:
1385:
1381:
1380:
1377:
1376:
1367:
1364:
1363:
1360:
1359:
1350:
1345:
1343:
1340:
1339:
1336:
1335:
1326:
1321:
1319:
1311:
1309:
1303:
1301:
1296:
1294:
1290:
1289:
1286:
1285:
1281:
1280:
1277:
1276:
1272:
1271:
1268:
1267:
1263:Leiosporoceros
1258:
1252:
1250:
1244:
1242:
1236:
1234:
1229:
1227:
1226:
1225:
1224:
1223:
1222:
1221:
1217:Phaeomegaceros
1213:
1205:
1197:
1176:
1175:
1174:
1173:
1172:
1171:
1150:
1149:
1148:
1147:
1146:
1145:
1137:
1129:
1121:
1113:
1096:Notothyladales
1092:
1091:
1090:
1089:
1088:
1087:
1079:
1071:
1054:Anthocerotales
1045:
1044:
1043:
1042:
1041:
1040:
1036:Leiosporoceros
1006:
1003:
948:Anthocerophyta
916:
915:Classification
913:
908:
900:
895:
844:
843:
823:
821:
810:
807:
799:pseudo-elaters
715:photosynthetic
664:
661:
633:pseudo-elaters
618:Early Devonian
555:grows from an
509:photosynthetic
485:
420:
417:
274:
273:
267:
266:
260:
259:
254:Classification
249:
248:
243:
241:Notothyladales
238:
233:
231:Anthocerotales
227:
226:
222:
221:
215:
214:
208:
207:
203:
202:
193:
189:
188:
183:
176:
175:
170:
166:
165:
152:
151:
137:
136:
128:
127:
111:
106:
101:
96:
91:
86:
81:
76:
71:
66:
61:
56:
55:
46:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
5635:
5624:
5621:
5619:
5616:
5614:
5611:
5609:
5606:
5605:
5603:
5586:
5581:
5577:
5573:
5567:
5563:
5558:
5554:
5550:
5545:
5541:
5537:
5531:
5527:
5522:
5518:
5514:
5509:
5505:
5501:
5496:
5492:
5488:
5483:
5479:
5475:
5470:
5466:
5462:
5457:
5453:
5449:
5444:
5440:
5436:
5430:
5426:
5421:
5417:
5412:
5406:
5402:
5397:
5391:
5387:
5386:
5384:
5382:
5378:
5374:
5369:
5355:
5354:
5349:
5347:
5346:
5341:
5331:
5328:
5326:
5323:
5321:
5318:
5316:
5313:
5311:
5308:
5306:
5303:
5301:
5298:
5296:
5293:
5291:
5288:
5286:
5283:
5281:
5278:
5276:
5273:
5271:
5268:
5266:
5263:
5261:
5258:
5256:
5253:
5251:
5248:
5246:
5243:
5241:
5238:
5236:
5233:
5231:
5228:
5226:
5223:
5221:
5218:
5216:
5213:
5211:
5208:
5206:
5203:
5201:
5198:
5196:
5193:
5191:
5188:
5186:
5183:
5181:
5178:
5176:
5173:
5171:
5168:
5167:
5165:
5163:
5159:
5153:
5150:
5148:
5145:
5143:
5140:
5138:
5135:
5133:
5130:
5128:
5125:
5123:
5120:
5118:
5115:
5113:
5110:
5108:
5105:
5103:
5100:
5098:
5095:
5093:
5090:
5088:
5085:
5084:
5082:
5080:
5076:
5070:
5069:Basidiomycota
5067:
5065:
5062:
5060:
5057:
5055:
5054:Glomeromycota
5052:
5050:
5047:
5045:
5042:
5040:
5037:
5036:
5034:
5032:
5028:
5022:
5019:
5017:
5016:Stramenopiles
5014:
5012:
5009:
5007:
5004:
5002:
4999:
4997:
4994:
4992:
4989:
4987:
4984:
4982:
4979:
4977:
4974:
4972:
4969:
4967:
4964:
4962:
4959:
4957:
4954:
4952:
4949:
4947:
4944:
4942:
4941:Apusomonadida
4939:
4937:
4934:
4932:
4929:
4927:
4924:
4923:
4921:
4918:
4913:
4910:
4908:
4904:
4894:
4891:
4889:
4886:
4884:
4881:
4878:
4877:Nanoarchaeota
4874:
4871:
4867:
4864:
4863:Euryarchaeota
4860:
4859:
4857:
4855:
4851:
4847:
4837:
4834:
4832:
4829:
4827:
4824:
4822:
4819:
4817:
4814:
4812:
4811:Spirochaetota
4809:
4807:
4806:Rhodothermota
4804:
4802:
4799:
4797:
4794:
4792:
4789:
4787:
4784:
4782:
4779:
4777:
4774:
4772:
4769:
4767:
4764:
4762:
4759:
4757:
4754:
4752:
4749:
4747:
4744:
4742:
4739:
4737:
4736:Dictyoglomota
4734:
4732:
4729:
4727:
4724:
4721:
4720:Cyanobacteria
4717:
4715:
4712:
4710:
4709:Chrysiogenota
4707:
4705:
4704:Chloroflexota
4702:
4700:
4697:
4695:
4692:
4690:
4687:
4685:
4684:Calditrichota
4682:
4680:
4679:Caldisericota
4677:
4675:
4672:
4670:
4667:
4665:
4662:
4660:
4657:
4655:
4654:Atribacterota
4652:
4650:
4647:
4645:
4642:
4640:
4637:
4635:
4632:
4631:
4629:
4627:
4623:
4619:
4615:
4610:
4602:
4597:
4595:
4590:
4588:
4583:
4582:
4579:
4567:
4564:
4562:
4554:
4552:
4548:
4545:
4541:
4540:
4537:
4489:
4486:
4484:
4481:
4479:
4476:
4475:
4474:
4471:
4469:
4466:
4465:
4463:
4457:
4453:
4447:
4444:
4442:
4439:
4437:
4435:
4430:
4428:
4425:
4424:
4422:
4416:
4412:
4406:
4402:
4400:
4396:
4394:
4390:
4388:
4384:
4382:
4378:
4376:
4372:
4370:
4366:
4364:
4360:
4358:
4354:
4352:
4348:
4346:
4342:
4340:
4336:
4334:
4330:
4328:
4327:Bennettitales
4324:
4323:
4321:
4308:
4307:Pteridosperms
4301:
4298:
4294:(seed plants)
4291:
4286:
4276:
4275:Protopityales
4272:
4270:
4266:
4264:
4260:
4258:
4254:
4253:
4251:
4248:
4241:
4237:
4234:
4228:
4217:
4213:
4210:
4208:
4204:
4202:
4198:
4197:
4195:
4191:
4187:
4181:
4177:
4175:
4171:
4170:
4167:
4164:
4161:
4156:
4145:
4141:
4137:
4133:
4130:
4128:
4124:
4122:
4118:
4117:
4115:
4113:
4109:
4105:
4102:
4096:
4086:
4082:
4080:
4076:
4074:
4070:
4069:
4067:
4060:
4056:
4053:
4046:
4045:Tracheophytes
4041:
4037:
4031:
4030:
4025:
4023:
4019:
4018:
4016:
4009:
4006:
4003:
3997:
3993:
3979:
3976:
3974:
3971:
3969:
3966:
3964:
3961:
3959:
3958:Andreaeopsida
3956:
3954:
3951:
3949:
3946:
3944:
3941:
3940:
3938:
3932:
3928:
3922:
3919:
3917:
3914:
3913:
3911:
3905:
3901:
3895:
3892:
3890:
3887:
3885:
3882:
3881:
3879:
3873:
3869:
3866:
3863:
3858:
3854:
3850:
3846:
3844:
3837:
3832:
3828:
3789:
3788:
3786:
3783:(land plants)
3780:
3776:
3770:
3767:
3766:
3764:
3760:
3754:
3751:
3749:
3746:
3745:
3742:
3739:
3737:
3733:
3727:
3724:
3722:
3721:
3717:
3715:
3712:
3710:
3707:
3706:
3704:
3701:
3696:
3692:
3676:
3675:Chlorophyceae
3673:
3671:
3668:
3666:
3663:
3662:
3661:
3658:
3656:
3653:
3651:
3650:Pedinophyceae
3648:
3646:
3643:
3641:
3638:
3636:
3633:
3632:
3630:
3628:Chlorophytina
3626:
3620:
3617:
3615:
3612:
3611:
3609:
3605:
3602:
3599:
3593:
3589:
3579:
3576:
3574:
3571:
3570:
3568:
3566:
3562:
3558:
3555:
3549:
3544:
3542:
3533:
3532:Viridiplantae
3529:
3519:
3516:
3515:
3513:
3511:
3507:
3501:
3498:
3497:
3495:
3493:
3489:
3483:
3480:
3478:
3477:Bangiophyceae
3475:
3473:
3470:
3468:
3465:
3463:
3460:
3458:
3455:
3453:
3450:
3449:
3447:
3441:
3437:
3431:
3430:
3426:
3425:
3423:
3421:
3417:
3411:
3410:
3406:
3405:
3403:
3401:
3397:
3393:
3390:
3387:
3382:
3378:
3366:
3363:
3361:
3358:
3356:
3353:
3352:
3351:
3348:
3346:
3343:
3342:
3341:
3338:
3334:
3331:
3329:
3328:Stramenopiles
3326:
3324:
3321:
3319:
3316:
3312:
3309:
3308:
3307:
3304:
3303:
3302:
3299:
3297:
3294:
3291:(major groups
3289:
3287:
3284:
3282:
3279:
3277:
3274:
3271:
3267:
3266:
3262:
3258:
3256:
3249:
3241:
3236:
3234:
3229:
3227:
3222:
3221:
3218:
3210:
3208:0-09-109301-5
3204:
3199:
3198:
3191:
3187:
3182:
3178:
3174:
3169:
3165:
3160:
3157:
3156:0-521-66097-1
3153:
3149:
3145:
3141:
3137:
3133:
3128:
3124:
3120:
3115:
3111:
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3102:
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3085:
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2713:
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2705:
2704:Nature Plants
2701:
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2657:
2653:
2649:
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2640:Nature Plants
2637:
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2612:
2607:
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2599:
2595:
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2562:1-56098-730-8
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2478:0-470-21359-0
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2196:
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2172:
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2162:
2157:
2151:
2148:. CRC Press.
2147:
2146:
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2018:
2013:
2008:
2003:
1999:
1995:
1991:
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1983:
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1156:
1155:
1152:
1151:
1143:
1142:
1138:
1135:
1134:
1133:Hattorioceros
1130:
1128:(1-2 species)
1127:
1126:
1122:
1119:
1118:
1114:
1111:
1110:
1106:
1105:
1104:
1103:
1099:
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966:
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953:
949:
945:
941:
937:
929:
927:
923:The hornwort
921:
914:
912:
906:
893:
889:
885:
881:
877:
873:
872:Andreaeopsida
869:
865:
861:
860:
855:
851:
840:
831:
827:
824:This section
822:
819:
815:
814:
808:
806:
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796:
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788:
783:
781:
777:
773:
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762:
758:
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749:
747:
743:
739:
735:
727:
722:
718:
716:
712:
711:multicellular
708:
704:
700:
696:
692:
687:
683:
679:
675:
670:
662:
660:
658:
654:
650:
646:
642:
638:
634:
630:
626:
621:
619:
615:
614:Late Silurian
611:
607:
602:
598:
597:
592:
591:
586:
585:
580:
579:
574:
570:
566:
562:
558:
554:
549:
547:
543:
539:
535:
534:
529:
528:
523:
522:cyanobacteria
519:
518:
513:
512:cyanobacteria
510:
506:
501:
499:
495:
491:
483:
479:
475:
471:
467:
466:
461:
460:
455:
454:
449:
445:
441:
437:
433:
429:
426:
418:
416:
413:
411:
410:
405:
399:
397:
393:
389:
385:
381:
375:
291:
287:
284:
280:
272:
271:Anthocerotae
268:
265:
261:
258:
256:
255:
247:
244:
242:
239:
237:
234:
232:
229:
228:
224:
223:
220:
217:
216:
212:
211:
209:
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197:
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104:
99:
94:
89:
84:
79:
74:
69:
64:
59:
53:
42:
39:
35:
34:
33:Ceratophyllum
19:
5380:
5351:
5343:
5265:Orthonectida
5260:Gastrotricha
5235:Nematomorpha
5210:Chaetognatha
5200:Hemichordate
5127:Pteridophyta
4996:Malawimonada
4870:Korarchaeota
4831:Thermotogota
4816:Synergistota
4791:Nitrospirota
4786:Nitrospinota
4731:Deinococcota
4664:Bacteroidota
4544:paraphyletic
4456:Angiospermae
4433:
4432:Ginkgoales (
4405:Petriellales
4387:Medullosales
4314:(seed ferns)
4027:
3948:Sphagnopsida
3943:Takakiopsida
3920:
3903:
3876:(liverworts)
3842:
3836:Embryophytes
3790:(see below↓)
3748:Charophyceae
3718:
3700:Streptophyta
3552:land plants)
3540:
3509:
3427:
3420:Rhodelphidia
3407:
3350:Opisthokonta
3254:
3196:
3185:
3172:
3163:
3143:
3135:
3131:
3122:
3118:
3109:
3105:
3088:. Retrieved
3077:
3044:
3040:
3030:
3011:
3007:
2994:
2982:. Retrieved
2971:
2946:
2942:
2917:
2913:
2907:
2864:
2860:
2850:
2815:
2811:
2801:
2766:
2762:
2752:
2707:
2703:
2697:
2643:
2639:
2628:
2585:
2581:
2571:
2552:
2546:
2501:
2497:
2487:
2468:
2448:
2437:
2394:
2388:
2378:
2347:
2341:
2296:
2292:
2281:
2270:
2235:
2231:
2221:
2178:
2174:
2164:
2144:
2137:
2110:
2106:
2096:
2059:
2055:
2045:
2034:
1989:
1985:
1975:
1950:
1946:
1936:
1891:
1887:
1877:
1866:
1819:
1815:
1805:
1796:
1786:
1761:
1757:
1751:
1720:The current
1635:
1634:
1618:
1617:
1594:
1593:
1569:
1568:
1533:Phymatoceros
1531:
1530:
1472:
1471:
1455:
1454:
1370:
1369:
1353:
1352:
1329:
1328:
1261:
1260:
1215:
1207:
1199:
1196:(43 species)
1191:
1184:
1179:
1167:Phymatoceros
1165:
1158:
1153:
1139:
1131:
1123:
1115:
1112:(21 species)
1107:
1100:
1095:
1081:
1078:(17 species)
1073:
1065:
1058:
1053:
1047:
1046:
1034:
1027:
1022:
1016:
1015:
1008:
992:
977:
962:
959:Anthocerotae
958:
956:
951:
947:
943:
940:paraphyletic
933:
924:
868:Sphagnopsida
859:Horneophyton
857:
847:
834:
830:adding to it
825:
784:
780:longitudinal
769:
750:
731:
725:
690:
685:
681:
677:
673:
666:
639:. They have
622:
600:
594:
588:
582:
576:
575:, the genus
572:
550:
537:
531:
525:
515:
502:
463:
457:
451:
422:
414:
407:
404:sub-tropical
400:
289:
286:Embryophytes
283:non-vascular
278:
277:
270:
251:
250:
195:
186:Embryophytes
179:
140:
38:
31:
5495:iNaturalist
5405:Wikispecies
5320:Brachiopoda
5295:Cycliophora
5240:Onychophora
5215:Kinorhyncha
5137:Ginkgophyta
5132:Cycadophyta
5097:Chlorophyta
5087:Glaucophyta
4781:Myxococcota
4699:Chlorobiota
4694:Chlamydiota
4351:Cordaitales
4345:Caytoniales
4231:Lignophytes
4214:(ferns and
4140:spikemosses
4079:Rhyniopsida
4029:Aglaophyton
3908:(hornworts)
3779:Embryophyta
3762:Anydrophyta
3720:Spirotaenia
3665:Ulvophyceae
3598:Chlorophyta
3548:green algae
3518:Algospongia
3492:Glaucophyta
3444:(red algae)
3014:: 150–166.
1953:: 181–188.
1737:Embryophyte
1637:Dendroceros
1220:(7 species)
1212:(7 species)
1204:(8 species)
1193:Dendroceros
1170:(2 species)
1144:(2 species)
1136:(1 species)
1086:(2 species)
1039:(1 species)
926:Dendroceros
850:crown group
746:chlorophyll
734:gametophyte
686:Dendroceros
596:Dendroceros
557:archegonium
538:slime pores
444:chloroplast
428:gametophyte
419:Description
409:Dendroceros
406:species of
396:gametophyte
392:gametophyte
5613:Bryophytes
5602:Categories
5353:Parakaryon
5315:Entoprocta
5250:Arthropoda
5245:Tardigrada
5225:Priapulida
5220:Loricifera
5205:Echinoderm
5175:Ctenophora
5147:Gnetophyta
5102:Charophyta
5092:Rhodophyta
5064:Ascomycota
5059:Zygomycota
5001:Metamonada
4976:Euglenozoa
4961:Ciliophora
4669:Balneolota
4644:Aquificota
4478:Magnoliids
4446:Gnetopsida
4216:horsetails
4144:quillworts
4136:clubmosses
4112:Lycophytes
4085:Renaliales
3862:Bryophytes
3440:Rhodophyta
3429:Rhodelphis
3090:7 December
2984:6 December
2698:Anthoceros
2107:BioScience
1743:References
1596:Nothoceros
1474:Phaeoceros
1457:Notothylas
1372:Anthoceros
1331:Folioceros
1209:Nothoceros
1117:Phaeoceros
1109:Notothylas
1075:Folioceros
1067:Anthoceros
999:antheridia
936:bryophytes
888:liverworts
884:bryophytes
864:sporangium
854:Cretaceous
787:sporophyte
765:antheridia
761:archegonia
753:monoecious
726:Phaeoceros
682:Nothoceros
663:Life cycle
645:bryophytes
637:liverworts
601:Notothylas
590:Nothoceros
578:Notothylas
565:liverworts
553:sporophyte
533:Folioceros
498:flavonoids
478:organelles
465:Anthoceros
459:Nothoceros
388:liverworts
380:sporophyte
192:Division:
120:Cretaceous
50:90–0
5608:Hornworts
5270:Dicyemida
5142:Pinophyta
5021:Telonemia
4956:Cryptista
4946:Breviatea
4931:Amoebozoa
4926:Alveolata
4907:Eukaryote
4659:Bacillota
3978:Bryopsida
3931:Bryophyta
3660:UTC clade
3409:Picomonas
3345:Amoebozoa
3323:Alveolata
3311:Cryptista
3286:Eukaryota
3069:209591279
3008:Phytotaxa
2421:0002-9122
2195:0032-0889
1722:phylogeny
1620:Megaceros
1201:Megaceros
1141:Mesoceros
1005:Phylogeny
880:Setaphyta
837:June 2008
757:dioecious
713:and even
703:protonema
691:germ tube
678:Megaceros
625:columella
584:Megaceros
527:Megaceros
453:Megaceros
279:Hornworts
169:Kingdom:
5562:35207907
5557:Tropicos
5432:BioLib:
5390:Wikidata
5330:Annelida
5325:Mollusca
5305:Phoronid
5300:Nemertea
5275:Rotifera
5255:Flatworm
5230:Nematoda
5195:Chordate
5185:Cnidaria
5180:Placozoa
5011:Rhizaria
4986:Haptista
4966:Cercozoa
4626:Bacteria
4488:Eudicots
4483:Monocots
4201:Ibykales
3935:(mosses)
3840:Plantae
3538:Plantae
3340:Amorphea
3318:Rhizaria
3306:Hacrobia
3296:Excavata
3281:Bacteria
3252:Plantae
3138:: 31–90.
3125:: 21–34.
2963:85582943
2920:: 7–16,
2899:53240320
2891:30411803
2842:33871608
2793:32790880
2744:32170292
2678:32042158
2620:36175544
2538:17030812
2429:21652320
2352:Elsevier
2333:17030812
2262:28584065
2213:35771646
2088:32117358
2026:23115334
1967:27194106
1928:23115334
1858:35360315
1731:See also
995:pyrenoid
980:Devonian
791:meristem
653:proximal
561:meristem
505:mucilage
474:pyrenoid
470:pyrenoid
382:. As in
264:Synonyms
44:Hornwort
5396:Q191156
5310:Bryozoa
5006:Provora
4981:Jakobea
4971:Discoba
4917:Protist
4854:Archaea
4607:Extant
4551:extinct
3400:Picozoa
3355:Animals
3276:Archaea
3112:: 1–62.
3049:Bibcode
2833:8321542
2784:7881058
2735:8075897
2712:Bibcode
2669:7027989
2648:Bibcode
2611:9630106
2590:Bibcode
2529:1622854
2506:Bibcode
2324:1622854
2301:Bibcode
2253:5462037
2204:9434304
2129:1313353
2079:7010833
2017:3503201
1994:Bibcode
1919:3503201
1896:Bibcode
1849:8964177
1824:bioRxiv
1778:3242017
984:species
928:crispus
905:RuBisCo
876:stomata
795:capsule
742:thallus
738:rosette
699:rhizoid
669:haploid
641:helical
569:stomata
546:stomata
542:thallus
482:RuBisCO
440:plastid
436:thallus
432:rosette
425:haploid
173:Plantae
124:present
5585:886300
5572:446447
5569:uBio:
5549:738980
5533:NZOR:
5513:500000
5425:238892
5170:Sponge
5162:Animal
4546:groups
4434:Ginkgo
4142:&
4000:
3596:
3333:Plants
3270:Domain
3205:
3154:
3067:
2980:. 2021
2961:
2897:
2889:
2840:
2830:
2791:
2781:
2742:
2732:
2676:
2666:
2618:
2608:
2559:
2536:
2526:
2475:
2427:
2419:
2397:(10).
2366:
2331:
2321:
2260:
2250:
2211:
2201:
2193:
2152:
2127:
2086:
2076:
2024:
2014:
1965:
1926:
1916:
1856:
1846:
1826:
1776:
988:genera
973:family
969:orders
892:mosses
803:spores
776:zygote
657:distal
629:tissue
517:Nostoc
490:plants
384:mosses
149:Prosk.
118:Upper
5580:WoRMS
5526:13809
5500:56327
5474:1AHOP
5448:622B3
5435:14876
5079:Plant
5031:Fungi
4951:CRuMs
3550:&
3360:Fungi
3065:S2CID
3004:(PDF)
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