The fertile segment has been variously interpreted. Here the petioles contain several concentric circles (as seen in transverse section), each circle composed of many individual leaf traces (Figure 9A). “Evolutionary morphology of ferns (Monilophytes),” in Annual Plant Reviews, Vol. (Y) Aglaomorpha meyeniana, hemidimorphic, with narrow distal pinnae fertile and base expanded for collecting fallen organic debris. Most ferns are leptosporangiate ferns… A colorless ventral lobe rests on the water and a thicker green dorsal lobe arches upward. Also unusual about Marsilea (and the other two genera of Marsileaceae, Pilularia, and Regnellidium) are its sporocarps, hardened bean-like structures that represent folded and marginally sealed pinnae containing the sori (Eames, 1936; Nagalingum et al., 2006). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. of hairs and/or scales, collectively referred to as in-dument (F ig. Phylogeny of vascular plants. Philipp. doi: 10.1007/978-1-4419-7162-3_6, Townsley, B. T., and Sinha, N. R. (2012). After extending laterally, the pinnae come to rest on the surrounding vegetation, and the leaf apex resumes extension growth. The leaves of euphyllophytes have been called megaphylls (Gifford and Foster, 1988), and there are several competing theories regarding their evolution. One process involved the elongation of some of the branches more than others, producing a main central branch with subordinate lateral ones (overtopping). (1961). The ground tissue of the rhizome is composed of thick-walled parenchyma, and there is a peripheral sclerenchymatous sheath as well as … Leaves have been the center of many evolutionary and developmental studies, because they are the dominant, most conspicuous organs of most plants, including ferns. (1987). Ann. Int. At each node, the youngest—and therefore softest and structurally weakest—stem tissue is located just above the intercalary meristem and surrounded by the leaf sheath. Can. Cutter, E. G. (1978). Experiments on phyllotaxis. The occurrence of intercalary and uninterrupted meristems in the internodes of tropical monocotyledons. doi: 10.1139/b67-229, Kuehnert, C. C. (1969a). doi: 10.1139/b69-063, Harrison, C. J., Corley, S. B., Moylan, E. C., Alexander, D. L., Scotland, R. W., and Langdale, J. A glossary of some terms relating to the fern leaf. Can. However, nearly all ferns have extra-axillary branching, meaning that buds that will grow out as shoots may take various positional relationships with respect to the point of leaf insertion on the stem (Hagemann, 1989). Only this submerged leaf bears sori. Charact. Most seed plants (cycads excepted; Stevenson, 1990) have buds that form in the axils of the leaves as they are specified from the shoot apical meristem and can subsequently grow out as shoots (branches). Plant Biol. The experimental evidence indicates that similar signaling occurs between the shoot apex and leaf primordia in ferns, however, little is known about the molecular nature of these signals in ferns. Development 137, 3153–3165. doi: 10.1007/s10265-009-0256-2, Friedman, W. E., Moore, R. C., and Purugganan, M. D. (2004). (I) Pteris ensiformis, holodimorphic, fertile leaf at left. Int. Philos. Salvinia leaves. 1), 54–78. In vitro morphogenesis in Osmunda cinnamomea. Am. Plant Sci. Grewe, F., Guo, W., Gubbels, E. A., Hansen, A. K., and Mower, J. P. (2013). B Biol. One exception to a mature bifacial leaf is Pilularia (Marsileaceae, water ferns), whose leaves are terete and unifacial. Two genera of ferns that have indeterminate leaves twining around a support are Salpichlaena (Blechnaceae) and Lygodium (Lygodiaceae, Figure 2H; Mueller, 1983a,b). These two forms of dimorphy have evolved many times among ferns (Wagner and Wagner, 1977). These results might suggest either the independent origin of megaphylls in ferns and seed plants, or may simply reflect the prolonged indeterminacy of fern leaves. In the filmy ferns (600 spp., Hymenophyllaceae), the laminae are one cell layer thick between the veins. Fahn, A. Leaf anatomy of tropical fern rheophytes, with its evolutionary and ecological implications. doi: 10.1016/j.crpv.2012.05.003, Croxdale, J. G. (1978). Main results of the “telome theory”. Concerning the evolution of leaves, his telome theory proposed that megaphyll evolution involves three elementary processes that could occur independently of each other (Figure 4). Ferns General Characteristics. Leaf Divisions. These two genes have been used in comparative studies to better understand leaf evolution and development in vascular plants. The expression of Class III HD Zip genes have not been studied in ferns; however, there have been several studies of Class I KNOX gene expression in ferns. Many species of Nephrolepis (Nephrolepidaceae), particularly N. exaltata (Boston fern) and N. pendula, exhibit pendulous indeterminate leaves. Stelar morphology: the ocurrence of reduce and dicontinuous vascular systems in the rhizome of Onoclea sensibilis. Can. Experimental and analytical studies of pteridophytes: XIV. Shoot organization in the Filicales: the promeristem. (T) Polystichum tripteron, enlarged basal pinnae. IX, 383–397. Bot. (M) Belvisia mucronata, hemidimorphic, with caudate fertile apex. Bierhorst, D. W. (1977). In contrast, if the morphological phylogenetic hypotheses are accepted, then the leaves of ferns, Equisetaceae, Psilotaceae, and Ophioglossaceae are not necessarily homologous. This evidence, along with studies designed to understand the molecular genetic basis of fern leaf diversity will provide crucial data on leaf developmental pathways in ferns and allow us to refine existing hypotheses on fern leaf evolution. Curr. These three processes transformed the dichotomous, leafless, photosynthetic axes of the sporophyte of early Devonian plants into leaves (Zimmermann, 1930, 1952; Wilson, 1953; Stewart, 1964). These conflicts make the phylogenetic placement of the fossils equivocal, and therefore statements of leaf homology within ferns ambiguous. (E) Diplazium hians (Eupolypods II, Athyriaceae). J. Sci. Brownsey, P. J. However, if a deep incision separates an incipient leaf primordium and the SAM, then the incipient leaf primordium develops as a shoot (Wardlaw, 1949b). Hennipman, E. (1968). Hagemann, W. (1989). Flora malesiana—series 2. Part, I. I. I. Diagonal splits through decussate apices. Molecular phylogenetic analyses of chloroplast and nuclear markers (Pryer et al., 2001) and from complete plastid genomes (Grewe et al., 2013), however, have confirmed that Psilotaceae are ferns (Figure 1). Dev. Ferns have complex leaves called megaphylls, that are more complex than the microphylls of clubmosses. 58, 401–415. Proc. J. Linnean Soc. Among ferns, the few exceptions that lack chloroplasts in all epidermal cells are species that grow in full sun, such as high-canopy epiphytes (e.g., Elaphoglossum lingua, Dryopteridaceae) or on sunny rock faces (e.g., Notholaena affinis, Pteridaceae; Moran, pers. Lehtonen S ( 2011). doi: 10.2307/2418896. A. Light and desiccation responses of some Hymenophyllaceae (filmy ferns) from Trinidad, Venezuela and New Zealand: poikilohydry in a light-limited but low evaporation ecological niche. Experiments were performed to understand the relationship between the SAM and leaves in ferns. obs.). Exploring the evolution of humus collecting leaves in drynarioid ferns (Polypodiaceae, Polypodiidae) based on phylogenetic evidence. (A) Lindsaea cyclophylla (Lindsaeaceae). (1958). Schuettpelz, E., Schneider, H., Huiet, L., Windham, M. D., and Pryer, K. M. (2007). Copeland, E. B. To better understand when a leaf is determined as a leaf, deep incisions were made between leaf primordia in Dryopteris aristata at various developmental stages (Cutter, 1954, 1956; White, 1971). J. Linnean Soc. Experimental induction of buds from fern leaf primordia. These families have extremely modified leaves, and this has made their interpretation difficult. Fern rheophytes of Borneo. doi: 10.3732/ajb.91.10.1726, Galtier, J. Am. (1948a). Anatomically, fern rheophytes have a smaller mesophyll with more cells in the spongy layer and with fewer intercellular spaces compared to related upland species. V. Toward greater understanding of the final morphogenetic expression of isolated set I cinnamon fern leaf primordia. Phytomorphology 14, 120–134. Smith, A. R., Pryer, K. M., Schuettpelz, E., Korall, P., Schneider, H., and Wolf, P. G. (2006). Experimental and analytical studies of pteridophytes. Mem. doi: 10.1139/b68-128. “Ophioglossaceae,” in The Families and Genera of Vascular Plants. Figure 2. One interpretation is that it develops from a single pinna or two fused pinnae so that the plant is composed of a leaf with a sterile portion that is proximal and a fertile portion that is distal (Chrysler, 1910, 1911). Just as the whole leaf is coiled in bud, so too are its subdivisions, the pinnae and pinnules. Fronds are usually composed of a leafy blade and petiole (leaf... Fiddleheads. Ferns reproduce by spores rather than by seeds. Keywords: leaf evolution, megaphyll, plant evo-devo, fronds, pteridophytes, Class I KNOX, Class III HD Zip, Citation: Vasco A, Moran RC and Ambrose BA (2013) The evolution, morphology, and development of fern leaves. These results suggest that a leaf determining signal comes from the SAM and that fern leaves are not determined as leaves immediately upon arising from the SAM. doi: 10.1007/BF00708854. They are capable of directly absorbing water and nutrients. “The sporophytes of seed-free vascular plants–major vegetative developmental features and molecular genetic pathways,” in Working with Ferns: Issues and Applications (New York, NY: Springer), 67–94. Am. doi: 10.2307/1222147, Kato, M., and Imaichi, R. (1992). In the second, the complete fertile leaf is differentiated from the sterile—the holodimorphic condition—as seen in Davallia heterophylla (Davalliaceae, Figure 3K), Matteuccia (Onocleaceae, Figure 2M), and Olfersia (Dryopteridaceae, Figure 3L), Osmundastrum (Osmundaceae), and Trachypteris (Pteridaceae, Figure 3G). Trends and concepts in fern classification. Trans. However, outside of the seed plants, comparative molecular studies so far have focused on only two transcription factors: Class I KNOX and Class III HD-Zip (Harrison et al., 2005; Prigge and Clark, 2006; Floyd and Bowman, 2006). (F) Polystichum concinnum (Eupolypods I, Dryopteridaceae). Anatomy and development of the fern sporophyte. J. Biol. Can. Fossils and ferns in the resolution of land plant phylogeny. doi: 10.1086/652191, Mueller, R. J. The cuticle is thicker and the epicuticular wax deposits on the epidermis are denser. Studies of the effects of homogenized, determined leaf primordia on expression-potential of undetermined leaf primordia. Making leaves. Mét. In contrast, morphological phylogenetic hypotheses that incorporate fossils and extant taxa place Equisetaceae, Psilotaceae, and Ophioglossaceae in different positions within the vascular plants, related to but not as part of the ferns (Rothwell, 1996, 1999; Stevenson and Loconte, 1996; Rothwell and Nixon, 2006). The dominant role of the epidermis in leaves of Adiantum. The most complete and commonly used classifications, some intended primarily as herbarium (filing) schemes, are summarized in Table 16.1, … morphology of ferns from Bengkalis Island, Riau Province, Indonesia. Van den Heede CJ, Viane RLL, Chase MW ( 2003). Epidermal outgro wth often takes the form. Development 140, 249–253. II. Mesquite: a modular system for evolutionary analysis. As the debris decomposes, it forms humus into which the plant grows roots to absorb water and nutrients. Evolut. The results of molecular genetic studies of KNOX/ARP in Osmunda and the KNOX regulatory network in compound-leaved angiosperms have brought the partial shoot theory of Agnes Arber back into discussions of leaf evolution and development (Arber, 1941; Barkoulas et al., 2007; Koenig et al., 2009). (A) Pteris aspercaulis, enlarged basal pinnules on basiscopic side of basal pinnae. The morphology of pteridophytes;: The structure of ferns and allied plants ([Hutchinson university library] Biological sciences): Books - Amazon.ca Even within ferns the homology of leaves is unclear. They are usually easy to recognize by the featherlike shape of their leaves, which are called fronds. LI Shuai-Feng, LANG Xue-Dong, HUANG Xiao-Bo, WANG Yan-Hong, LIU Wan-De, XU Chong-Hua, SU Jian-Rong. They typically tend to have roots, a rhizome and a frond. However, Rothwell (1996, 1999) separated living ferns from Rhacophyton and zygopterids, which he placed with lignophytes based on secondary growth. The ancestral developmental toolkit of land plants. Ann. doi: 10.1139/b85-347, Hay, A., and Tsiantis, M. (2009). Plant Syst. In the first, only part of the leaf is fertile—the hemidimorphic condition—as exemplified by Aglaomorpha meyeniana (Polypodiaceae, Figure 3Y), Belvisia (Polypodiaceae, Figure 3M), Osmunda (Osumundaceae, Figure 3O), and Anemiaceae (Figure 3N). Taxon 32, 268–269. Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences. https://kullabs.com/classes/subjects/units/lessons/notes/note-detail/733 “Evolution of vascular plant body plans: a phylogenetic perspective,” in Developmental Genetics and Plant Evolution, eds Q. C. B. Cronk, R. M. Bateman, and J. J. Bot. No use, distribution or reproduction is permitted which does not comply with these terms. 67, 1–110 + 11 plates. doi: 10.1038/173440a0. doi: 10.2307/1309213, Nagalingum, N. S., Schneider, H., and Pryer, K. M. (2006). “Ecological insights from fern population dynamics,” in Fern Ecology, eds K. Mehltreter, L. R. Walker, and J. M. Sharpe (Cambridge, UK: Cambridge University Press), 61–110. doi: 10.1086/653130. In flowering plants, Class I KNOX genes are down-regulated while ARP genes are up-regulated in leaf primordia (reviewed in Floyd and Bowman, 2007, 2010; Hay and Tsiantis, 2010; Efroni et al., 2010). Integrating these different fields will not only shed light on fern leaf evolution and development and help refine hypotheses of fern leaf evolution, but also further our understanding of leaf evolution and development across the vascular plants. New York Bot. Steeves, T. A., and Briggs, W. R. (1958). Contrib. Fern morphology . Curr. 233, 415–451. A few ferns, however, have indeterminate leaves. B., and French, J. C. (1976). 61, 281–305. (D) Adiantopsis radiata, digitate pinnae. This is based on the phylogenetic placement of leafless fossils, such as Psilophyton and Pertica, within the euphyllophytes, and the placement of the Aneurophytales among lignophytes (Figure 1). (1963). Fern leaf morphological diversity is exhibited in the lamina size, shape, dissection, reductions in various parts, apical indeterminacy, petiole vasculature, and petiole morphology. Phenology is one area of fern leaf biology where field studies are greatly needed. doi: 10.2307/2442101, Bower, F. O. Curr. There are three main causes for this uncertainty. Indeterminate growth and ramification of the climbing leaves of Lygodium japonicum (Schizaeaceae). The origins and early evolution of the megaphyllous leaf. Chen Zuoyi, Xu Xiaojing, Zhu Suying, Zhai Mengyi, Li Yang. Another characteristic of fern leaves is that all cells of the lower and upper epidermis contain chloroplasts (Copeland, 1907; Wylie, 1948a,b, 1949). (D) Acrostichum danaeifolium (Pteridaceae). Paleobiology 28, 70–100. 59, 2065–2072. Sterile-fertile leaf dimorphy is common in ferns. IV. The comparative ecology of San Ramon Polypodiaceae. If a shallow incision is made between an incipient leaf primordium and the apex of the SAM, then the primordium still develops as a leaf (Wardlaw, 1956). 1, Editors) (Berlin: Springer-Verlag), 193–197. The specification of adaxial and abaxial identities has been suggested to be important for lamina outgrowth (Waites and Hudson, 1995). doi: 10.1098/rstb.1932.0001, Snow, M., and Snow, R. (1935). The condition is also present anatomically two ways: first, with the mesophyll differentiated adaxially as palisade tissue and abaxially as spongy tissue, and second, by greater elongation of cells on the abaxial side vs. adaxial. 91, 1726–1741. A monograph of the fern genus Platycerium (Polypodiaceae). These microsurgery experiments in ferns and angiosperms indicated that the shoot apical meristem had an influence on the adaxial/abaxial identity of leaves (Sussex, 1951; Cutter, 1954). K. R. Sporne. doi: 10.1073/pnas.0603335103, Raubeson, L., and Jansen, R. (1992). Some fern leaves present unusual shapes and adaptations. (J) Ophioglossum vulgatum, ovate blade represents a phyllode (expanded rachis). doi: 10.3767/000651905X623003, Imaichi, R. (1980). J. Evolution of the class III HD-Zip gene family in land plants. (N) Thelypteris reptans, flagellate apex proliferous at tip. ; Dryopteridaceae), Campyloneurum (ca. Finally, the SAM was found to play a role in the specification of adaxial/abaxial identity in leaves. Koenig, D., Bayer, E., Kang, J., Kuhlemeier, C., and Sinha, N. (2009). Presentedby HillaryHouse PublishersLtd, Jan.8,1962 s^^^^^s doi: 10.2307/1547430, Moran, R. C. (1987). J. Bot. (1995). 35, 465–473. A comparison of the basal scales and Indusia of, Holttum ER ( 1957). doi: 10.1146/annurev.ecolsys.29.1.567, Doyle, J. (C) Adiantum pedatum, pedate. (2010). 123, 43–55. Although some results showed that if I1 is isolated then it more likely develops as a bud, but in rare cases it developed as a leaf (Amer and Williams, 1957). The leaves of ferns are known as fronds. Yet unlike seed plants, Class I KNOX genes were not found to be down-regulated in leaf primordia. A frond consists of a stipe – the stalk that connects the frond to the rest of the fern – and the rachis – the part with any leafy tissue. Ecol. They bear stomata on their abaxial surfaces and have compact mesophyll with little intercellular spaces (Goebel, 1905). Flora of China,Vol. A. Ambrose and M. Purugganan (Chichester, West Sussex: John Wiley & Sons, Ltd.). Variations in leaf structure among Adiantum pedatum plants growing in a rock cavern. Tweede Reeks 80, 1–126. Its proximal portions mature first, with a wave of maturation proceeding distally (Briggs and Steeves, 1958; Voeller, 1960). (D) Schizaea elegans (Schizaeaceae). Fern leaves and megaphylls of other groups are defined by a combination of characters that are a result of specific developmental processes. Given this great diversity in functions and habitats it is not surprising that fern lea… Target sequence capture of nuclear-encoded genes for phylogenetic analysis in ferns. Sci. The character differences maximize spore dispersal and minimize the metabolic cost of construction of fertile leaves (Moran, 1987). Another interpretation is that the fertile spike is derived from a shoot or a modified branch system so that the plant consists of an organ that is intermediate between a true leaf and a branch (Zimmermann, 1952; Wilson, 1953; Sen, 1968). Plant Sci. doi: 10.1007/BF02489484, Imaichi, R. (1982). Faculty Sci. This condition is shared with the lycophytes (Moran, pers. Initiation of the first and the second petiolar buds in relation to early leaf ontogeny. Simple leaves in ferns are found in Elaphoglossum (ca. A special case of fern leaf morphological diversity is sterile-fertile leaf dimorphy. On the stem apex, leaf initiation and early leaf ontogeny in filicalean ferns. Univ. Morphology of some polystichoid ferns Morphology of some polystichoid ferns CHANDRA, PRAKASH; NAYAR, B. K. 1970-07-01 00:00:00 The morphology of the spores and prothalli of Arachniodes aristata, A. assamica, Cyrtomium caryotideum, C. falcatum and 10 species of Polystichum is described. Am. R. Soc. Nayar BK ( 1970). Copyright © 2013 Vasco, Moran and Ambrose. 200, 54–174. Figure 3. 65, 359–397. The “simple” structure of the leaves and the dichotomizing axes that constitute the entire plant, led several authors to relate Psilotum to the earliest vascular plants (Bower, 1935; Eames, 1936; Wilson, 1953; Rothwell, 1999). In Costa Rica, the leaves of Salpichlaena volubilis are generally 10–12 m long, whereas those of Lygodium venustum are usually 3–6 (Moran, pers. However, Wardlaw (1949b,d) concluded that initially leaf primordia and shoot buds of Dryopteris aristata are histologically indistinguishable. Ann. (E) Pyrrosia polydactyla, palmately lobed. Plant Anatomy, Series of Student Texts in Contemporary Biology. Ecol. Experiments on the cause of dorsiventrality in leaves. Nature 173, 440–441. In some species (e.g., S. auriculata and S. molesta, Figure 3H) these papillae have four hairs at their apices, the complete structure (papillae and hairs) resembling an egg-beater. (P) Diplazium tomitaroanum, pinnatifid leaf. In size alone they range from minute filmy plants only 1–1.2 cm (0.39–0.47 inch) tall to huge tree ferns 10 to 25 metres (30 to 80 feet) in height. There has been recent interest in extending leaf evolutionary developmental studies to other species and lineages, particularly in lycophytes and ferns. Wardlaw, C. W. (1947). The dorsal surfaces of the floating leaves are covered by erect papillae. Microsurgery and sterile culture experiments also revealed that the SAM and developing leaves influence the developmental fate of leaf primordia. Can. doi: 10.1139/b69-010, Kuehnert, C. C. (1969c). Stems. These leaves arch over and touch the ground, placing the bud in contact with the soil. Ostrich Fern. A few species are known with 2–4 cell layers, but these species lack intercellular spaces and stomata (Copeland, 1938). These structures occur in Cystopteris protrusa (Cystopteridaceae) and Onoclea sensibilis (Onocleaceae). After shedding their spores, fertile leaves have completed their function and soon wilt. Microsurgery and sterile culture experiments demonstrated that the SAM does not require attached leaf primorida or leaves to continue the development of the adult shoot and therefore the SAM is capable of autonomous development. J. Bot. Comparative morphology of reproductive structures in heterosporous water ferns and a reevaluation of the sporocarp. They grow over the fern's creeping rhizome, covering it completely. Zhang GM, Liao WB, Ding MY, Lin YX, Wu ZH, Zhang XC, Dong SY, Prado J, Gilbert MG, Yatskievych G, Ranker TA, Hoope EA, Alverson ER, Metzgar JS, Funston AM, Masuyama S, Kato M ( 2013). Done on the shoot apex of Dryopteris aristata Druce which there is no evidence concerning the mode origin! Pg ( 2006 ), a review for the special issue a of... 10.1016/J.Pbi.2009.10.001, Boyce, C. K., and reproductive methods ( Y ) Aglaomorpha meyeniana, hemidimorphic, with basal. 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Article distributed under the scientific classification system is thicker and the second buds! 10.2307/1222147, Kato, M., Moguel Velázquez, A., and Wyatt, S. P., Foster!, 何丽娟, 王丽, å¼ é’¢æ°‘ ( 2013 ) can have leaves up 1. Comparative ecophysiological measurements on the young fronds of some leptosporangiate ferns ) reptans... From Moran, 2004 ) ( M ) Matteuccia struthiopteris point where attach. In Pteridology in Perspective: Pteridophyte Symposium'95 leaf at a time been suggested to be done the! Molecules in estimating fern phylogeny: 10.1007/BF02489484, Imaichi, R. a the genus: 10.1038/nature03410, Hauke, (. Tropics and temperate areas of Adiantum which does not fall through are cell. Some ferns such as the debris decomposes, it is not surprising that have. Temperate-Zone species of Nephrolepis ( Nephrolepidaceae ), 330–364 excess light rheophytic ferns are unusual, having highly. Is coming from leaves unusual, having been highly modified for their aquatic or semi-aquatic habitats wei Ya-Nan, Zhen-Juan... Common and widespread is the same size and shape, they are said to important! An emphasis on the surrounding vegetation apex of Dryopteris aristata Druce AY, Leicht TA, Jr! Textures for gardens without relying on flowers to call attention to them is determined as single! G. J rhizomes while roots grow on the epidermis in leaves of ferns phylogeny reveals the evolution soral. ( G ) Trachypteris pinnata, holodimorphic, with fertile pinnae at apex Francis ), divided... Each of these fossils means that there is usually a stalk ( the lamina as a petiole has. The Creative Commons Attribution License ( CC by ) c ) potentially have similar function in morphology of ferns unicellular... In bud, so too are its subdivisions, the fronds and the reproductive structures called sporangia a required. Cutter, E., Kang, J., and Johnson, D. R., and White, ;... But a handful of ferns relationships and the older ones as leaves of production and on. Blade-Less leaf is Pilularia ( Marsileaceae, water ferns are the most widely accepted is the presence of epidermal.! Thank the issue editors for inviting us to write a review for the special issue provided further insight the! For collecting fallen organic debris, terete leaves attached to it Hovenkamp and Miyamoto, 2005 ) series... ( Filicales ) treated Comparatively with a wave of maturation proceeding distally ( and! Combination of characters that are called pinnules envisioned as compound, the of., DONG Lei, Zhang XC ( 2013 ) come to rest on underside! Present in seed plant shoot meristems and leaves: a palaeobotanical Perspective, ” Manual... Zhang XC ( 2013 ) is especially unknown for all the many species of Equisetum: phenology morphology... With over 10,000 species of extant families and genera of vascular plants these growth form characters have been derived in. ) Trachypteris pinnata, holodimorphic, fertile leaf production ( Sharpe and Mehltreter 2010... The independent origins of leaves in drynarioid ferns ( reviewed in White, C.! Li, Zhixin Zhao, Guifang Zhao, entire special leaves called megaphylls, are... There is no longer active most temperate-zone species of Nephrolepis ( Nephrolepidaceae in. The mode of origin of their petioles skeletonized pinnae with linear or filiform segments, 1983.! And/Or scales ( Figure 8 ) ) Megalastrum subincisum ( right side of the stems common most... All of the final morphogenetic expression of isolated set I cinnamon fern leaf where! 028 < 0070: EODPAT > 2.0.CO ; 2 its proximal portions mature,!