morphology of ferns

Read "Overview of the morphology, anatomy, and ontogeny of Adiantum capillus‐veneris: An experimental system to study the development of ferns, Journal of Systematics and Evolution" on DeepDyve, the largest online rental service for scholarly research with thousands of academic publications available at your fingertips. Contrib. Figure 2. The leaf is formed by divisions of the apical initial as well as by divisions in the marginal meristem. In highly divided leaves, branching patterns of the pinnae (primary divisions of the laminae) may be of taxonomic importance (e.g., species of Megalastrum (Figure 2I) and Lastreopsis, both Dryopteridaceae). This research was supported by a grant to Moran and Ambrose from the United States National Science Foundation (DEB-1020443). Experimental studies have shown that older leaves not only inhibit the growth and placement of nearby incipient leaf primordia but also have a role in determining whether a leaf primordium develops as a leaf or a shoot (reviewed in White, 1971). 47, 69–72. Bower, F. O. They bear stomata on their abaxial surfaces and have compact mesophyll with little intercellular spaces (Goebel, 1905). However, if a deep incision separates an incipient leaf primordium and the SAM, then the incipient leaf primordium develops as a shoot (Wardlaw, 1949b). Leaves are generally considered to be determinate organs; that is, they grow only to a certain length and no more. doi: 10.2307/1222147, Kato, M., and Imaichi, R. (1992). 维管植物比较形态学. Shoot organization in the Filicales: the promeristem. Epidermal outgro wth often takes the form. Morphology and Evolution of Vascular Plants, 3rd Edn. One of these genera of humus collectors, Aglaomorpha (Polypodiaceae, Figure 3Y), has sterile and fertile leaves of one type (monomorphic) but modified within the same leaf to accumulate organic debris. (H) Lygodium flexuosum, rachis (at right) with lamina of pinnule (other half of pinna not shown). Bot. Some examples of rheophytic ferns are Asplenium obtusifolium (Aspleniaceae), Osmunda lancea (Osmundaceae), and Tectaria lobbii (Tectariaceae). The Origin and Early Diversification of Land Plants: A Cladistic Study. obs.). In addition, isolated SAMs grown in sterile culture on media supplemented with sucrose and auxin formed adult plants (White, 1971). Washington, DC: Smithsonian Institution Press. That half of a pinna or pinnule that occurs on the side toward the distal apex of the axis that bears it is called the acroscopic side, and accordingly, the side that occurs toward the base is the basiscopic (for more details of fern leaf terminology, see Tryon, 1960). obs.). Isolated P3 leaves were also grown with older leaves that were not in physical contact with each other. Their surfaces bear stomata that allow air to diffuse into the loosely packed parenchyma beneath the line. Only the fern families mentioned in the text are included here, for a complete fern family phylogeny see Smith et al. If all of the leaf primordia are removed from the SAM, then the incipient leaf primordium (I1) develops as a leaf (Hicks and Steeves, 1969; White, 1971). In temperate species, the apical meristem differentiates as parenchyma by the end of the summer and therefore is no longer active. Ecol. Rai HS, Graham SW ( 2010). These studies found that, as in seed plants with compound leaves, Class I KNOX genes are expressed in the meristem and in the developing leaves. Figure 6. 35, 465–473. doi: 10.2307/1221976. Indeterminate growth of the climbing leaves of a fern. Can. The experimental induction of buds from leaf primordia in Dryopteris aristata Druce. Experiments on phyllotaxis. In Equisetum, the relationship of the vegetative and fertile portions has been debated: the sporangiophore (the fertile portion) has been interpreted as being a novel organ “(organ sui generis)” or homologous to the leaves (Goebel, 1905; Bower, 1935; Zimmermann, 1952; Page, 1972). Am. (M) Belvisia mucronata, hemidimorphic, with caudate fertile apex. 115-140. In some species of Diplazium (Athyriaceae) with erect subarborescent rhizomes (e.g., Diplazium prominulum and D. striatastrum), the rigid, starch-filled trophopods are tightly appressed to the trunk and structurally support the rhizome (Moran, pers. Am. Cambridge, England; New York: Cambridge University Press. The Ferns (Filicales), Vol. Philipp. (1926). They are capable of directly absorbing water and nutrients. Plant Biol. This is not a thigmotropic grasping response as exhibited, for instance, by the leaves of Clematis (Ranunculaceae; Darwin, 1876). 1, Editors) (Berlin: Springer-Verlag), 193–197. R. Soc. After shedding their spores, fertile leaves have completed their function and soon wilt. 57, 1951–1959. 57, 8–55. A frond is simply the leaf of the fern. Sci. doi: 10.2307/1547430, Moran, R. C. (1987). Lond. Aerophores or pneumatophores are also characteristic of nearly all leptosporangiate fern leaves. The SAM in ferns typically has 1 or 2 large apical cells surrounded by small cytoplasmically dense cells that divide frequently (Bower, 1884; Wardlaw, 1963; Bierhorst, 1971; White, 1971; McAlpin and White, 1974; Stevenson, 1976; White and Turner, 1995). Ann. (J) Ophioglossum vulgatum, ovate blade represents a phyllode (expanded rachis). Steeves, T. A. From genes to shape: regulatory interactions in leaf development. Bot. Scrambling ferns often have leaf apices exhibiting intermittent growth (Figure 8). Impact Factor 4.402 | CiteScore 7.8More on impact ›, Understanding plant adaptation and diversification: evo-devo and beyond (1948a). Proceedings of the Holttum Memorial Pteridophyte Symposium, eds J. M. Camus, M. Gibby, and R. J. Johns (Kew: Royal Botanic Gardens), 395–404. Growth 13, 93–131. doi: 10.1139/b81-268, Cutter, E. G. (1954). The principal attribute distinguishing Ophioglossaceae from other ferns is the division of the leaf into separate vegetative (sterile segment) and sporangium-bearing (fertile segment) portions (Figure 2J; Bower, 1926; Gifford and Foster, 1988; Wagner, 1990). Morphogenetic Studies on Osmunda cinnamomea L: The auxin relationships of expanding fronds. The Morphology of Pteridophytes. Fern leaf primordia arise from one or a group of cells on the flank of the SAM (Bower, 1923; Wardlaw, 1949b; Steeves and Briggs, 1958; Bierhorst, 1977; Imaichi, 1980, 1982). doi: 10.1086/653130. 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). Eventually the leaf comes to rest on the soil and the bud, still attached to the leaf, is “planted” and takes root. This fern is originated from the Eastern Continent. Ferns and fern relatives are known as Pteridophytes. A brief comparative survey of aerophore structure within the Filicopsida. 233, 415–451. (1995). This suggests that another part of the plant has a determining influence on leaf development besides older leaves. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. These controversies along with the phylogenetic position of ferns as sister to seed plants, and the fact that fern leaves display a great morphological diversity, make ferns a key plant lineage for comparative studies on how leaves and vascular plants evolved. 2. Mag. Proceedings of the Holttum Memorial Pteridophyte Symposium, eds J. M. Camus, M. Gibby, and R. J. Johns (Kew), 435–467. 22, 2325–2328. 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. (1983b). Development 136, 2997–3006. Most commonly, as the leaf senesces, the petiole weakens at the base, and the leaf gradually reclines toward the ground. New York, NY: D. Appleton and Company. Stevenson, D. (1990). Comparative morphology of reproductive structures in heterosporous water ferns and a reevaluation of the sporocarp. Developmental potentialities of leaf primordia of Osmunda cinnamomea. In angiosperms, polar auxin transport is important in specifying phyllotaxy, leaf patterning, and is integral in the leaf development network involving 2 key transcription factors, Class I KNOX and Class III HD-Zip (see below). Can. Further experimental observations on the shoot apex of Dryopteris aristata Druce. doi: 10.1126/science.1070343. Figure 5. 55, 223–231. This condition is shared with the lycophytes (Moran, pers. 167, 805–815. Wylie, R. B. Alansmia (Polypodiaceae) is a pendent epiphyte whose leaves exhibit small continuously growing fiddleheads at their tips (Kessler et al., 2011). Evolut. KNOX homeobox genes potentially have similar function in both diploid unicellular and multicellular meristems, but not in haploid meristems. A. Ambrose and M. D. Purugganan (John Wiley and Sons Ltd), 115–140. Phyllotaxis and organogenesis in ferns. (1989). The Movements and Habits of Climbing Plants, 2nd Edn., New York, NY: D. Appelton and Company. Kato, M. (1988). doi: 10.1093/jxb/11.1.45, Steeves, T. A., and Sussex, I. M. (1989). Sterile-fertile leaf dimorphy is common in ferns. (Y) Aglaomorpha meyeniana, hemidimorphic, with narrow distal pinnae fertile and base expanded for collecting fallen organic debris. Arber, A. These experiments indicate that the SAM is not only capable of autonomous development but also has a leaf-determining influence on incipient primordia. J. Bot. Leaves are lateral determinate structures formed in a predictable sequence (phyllotaxy) on the flanks of an indeterminate shoot apical meristem. Megaphylls then are present in seed plants and ferns and there are several competing theories regarding their evolution and origin. Opin. At the base of each leaf sheath is a node at which there is an intercalary meristem. Bot. doi: 10.1600/036364409789271209. Wagner, W. H. Jr. (1990). B., and French, J. C. (1976). The root system is matted and tightly anchored to the substrate. One example of simple leaves are the reniform ones that have evolved independently in different fern families, resulting in some striking parallelisms (Figure 7). (1995). If incisions were made close to I1 then the primordia that grew out were radially symmetric. The characteristics of each of these 3 parts of the fern plant are used for classification and identification. This prolonged meristematic activity is due to an apical cell at the leaf tip and is a distinctive feature of ferns compared to seed plants (Imaichi, 2008). Many species of Nephrolepis (Nephrolepidaceae), particularly N. exaltata (Boston fern) and N. pendula, exhibit pendulous indeterminate leaves. 91, 717–727. 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. Fronds are usually composed of a leafy blade and petiole (leaf... Fiddleheads. In Dryopteris aristata, if all of the surrounding primordia are removed from a growing shoot tip, then the apical meristem continues to grow and produce leaves (Wardlaw, 1947, 1949a,c; White, 1971). Darwin, C. (1896). Leaf anatomy of tropical fern rheophytes, with its evolutionary and ecological implications. Daigobo, S. (1972). Iowa Acad. (2011). Moran, R. C., Labiak, P. H., and Sundue, M. (2010). Cyatheaceae. How does the inclusion of fossil data change our conclusions about the phylogenetic history of euphyllophytes. FANG Jing-Yun, GUO Ke, WANG Guo-Hong, TANG Zhi-Yao, XIE Zong-Qiang, SHEN Ze-Hao, WANG Ren-Qing, QIANG Sheng, LIANG Cun-Zhu, DA Liang-Jun, YU Dan. A glossary of some terms relating to the fern leaf. (1938). There are two characteristics typical of most fern leaves: a fiddlehead and aerophore lines (Figure 6). V. Toward greater understanding of the final morphogenetic expression of isolated set I cinnamon fern leaf primordia. In addition, experimental studies found that fern leaves are not determined as leaves until later in their development but when first specified have more of a shoot identity (reviewed in White, 1971). Bot. If the molecular phylogenetic hypothesis is accepted, then all ferns leaves should be homologous at some level. Even within ferns the homology of leaves is unclear. Ostrich Fern. As an adaptation to their epiphytic life style, the drynarioid genera of Polypodiaceae have leaves modified for collecting organic debris that falls from above, mostly bits of bark and leaves (Hennipman and Roos, 1982; Janssen and Schneider, 2005). Phylogeny and character evolution of the bolbitidoid ferns (Dryopteridaceae). 600 spp. Received: 28 May 2013; Accepted: 15 August 2013; Published online: 04 September 2013. Its blade-less leaf is interpreted as a petiole that has lost its apical pinnae (Eames, 1936). Biol. “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. 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. This is important nutritionally because these ferns are epiphytic, not in contact with water and minerals in the soil. 99, 85–106. doi: 10.1139/b68-128. A phylogenetic classification of the homosporous ferns. In nearly all extant ferns, leaves constitute the dominant organs of the plant. Plant Res. Tryon RM, Tryon AF ( 1982). Pryer KM, Schuettpelz E, Wolf PG, Schneider H, Smith AR, Cranfill R ( 2004). Experimental and analytical studies of pteridophytes. The origin and evolution of leaves in vascular plants has been widely debated. The genus Paesia in Malaysia. 45. 59, 2065–2072. They are downregulated in determinate simple leaves but upregulated in compound leaves to specify pinnae (Bharathan et al., 2002; reviewed in Hay and Tsiantis, 2009). Morphogenetic studies of fern leaves. Mol. Further support for the shoot-like characteristics of fern leaves comes from recent studies in Nephrolepis exaltata (Nephrolepidaceae) that found that there is a reiteration from shoot apical meristem to leaf to pinnae that suggests a reiteration of a shoot program (Sanders et al., 2011). Evolut. |, The Morphological Diversity of Fern Leaves, Experimental Analyses of Fern Leaf Development, Molecular Genetics of Fern Leaf Development, Creative Commons Attribution License (CC BY), The New York Botanical Garden, Bronx, NY, USA. Experimental studies of the sporophytes of ferns. 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. Presentedby HillaryHouse PublishersLtd, Jan.8,1962 s^^^^^s 91, 1726–1741. Ecol. Furthermore, ferns grow in many habitats—from mangroves at sea level to alpine vegetation above tree line, temperate forests to arctic tundras, and desserts to wetlands. Articles. They grow tightly overlapped and appressed or ascending over rhizome, with sessile and broad bases—all modifications so that trapped organic matter does not fall through. Rev. Experiments on organogenesis in ferns. Torrey Bot. Plant Sci. The evolutionary patterns of living ferns. As new fronds emerge, generally in the spring, they unroll, these unrolling fronds are called fiddleheads. Rarely, foliar-borne buds abscise from the mature leaf and drop to the ground, as in Cystopteris bulbifera (Cystopteridaceae, Figure 3W). This study found that the expression of Class I KNOX and ARP genes overlapped in the meristem and leaves unlike the complementary expression patterns in flowering plants (Harrison et al., 2005). doi: 10.1111/j.1095-8339.1991.tb00220.x, Doyle, J. Leaf primordia P1, P2, and P3 are more plastic in their development and after incisions, may grow out as buds, although a few still develop as leaves. doi: 10.1086/600135, Sano, R., Juárez, C. M., Hass, B., Sakakibara, K., Ito, M., Banks, J. doi: 10.1002/adma.200904411, Bharathan, G., Goliber, T. E., Moore, C., Kessler, S., Pham, T., and Sinha, N. R. (2002). (1923). (E) Diplazium hians (Eupolypods II, Athyriaceae). Fern leaves and megaphylls of other groups are defined by a combination of characters that are a result of specific developmental processes. Hennipman, E. (1968). Phytomorphology 11, 346–359. Wardlaw, C. W. (1945). doi: 10.1093/aob/mcg077, Proctor, M. C. F. (2012). The field of developmental genetics is the missing piece and when integrated with data from other fields will help us develop more robust hypotheses of fern leaf evolution and development and more broadly hypotheses of leaf evolution and development in vascular plants. 12, 593–598. Hypotheses of megaphyll homology depend on the morphological and anatomical interpretations of extinct leafless Devonian and Carboniferous plants. In Dryopteris aristata, incisions were made to isolate incipient leaf primordia from the SAM and/or older primordia. Although fern leaves are often stereotyped as being finely divided, some are simple and entire, and others are merely lobed. 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. A community-derived classification for extant lycophytes and ferns. (N) Thelypteris reptans, flagellate apex proliferous at tip. Neerl. 3(Suppl. Trans. Evolutionary morphology of ferns (monilophytes). The twining results from the widely circumnutating leaf being interrupted by contact with a support, and, when the rope hits a pole, the rope continues its motion around the pole, wrapping upward and thus climbing the pole (Darwin, 1876). Sometimes the basal pinnae are repeatedly branched on the basiscopic side—a condition known as pedate. 62, 1336–1343. The cytohistological and cytohistochemical zonation of the shoot apex of Botrychium multifidum. The Nephrolepis pendula complex (Lomariopsidaceae) in the Neotropics. These conflicts make the phylogenetic placement of the fossils equivocal, and therefore statements of leaf homology within ferns ambiguous. Interestingly, these are exactly the roles played by Class I KNOX and Class III HD-Zip genes, respectively. Although the former species is now classified in Arachniodes (Dryopteridaceae) and the latter in Osmundastrum (Osmundaceae), we will use here the older names employed by the researchers in the original developmental studies cited. Brownsey, P. J. Experiments were performed to understand the relationship between the SAM and leaves in ferns. J. Maturation of the developing fern leaf is acroscopic, that is, toward the apex (Figure 5). These lines aerate the leaf. II. Fossil evidence has also allowed some authors to hypothesize that the full set of megaphyllous traits were acquired later, either at the same (Boyce and Knoll, 2002) or in a different (Sanders et al., 2009) sequence of events in ferns compared to seed plants. Fern leaves generally exhibit finite (determinate) growth but with longer meristematic activity of the apical portion and maturation toward the apex (acroscopic growth). Taxon 37, 381–386. London: MacMillan and Co. Boyce, C. K. (2010). Through the action of the pulvini, the pinnae fold forward and upward at night, forming a vertical packet with the basal pair enclosed within the distal one. They grow over the fern's creeping rhizome, covering it completely. Paleobotanical evidence supports the existence of these different processes and also suggests that there were anatomical changes that correlated with the origin of megaphylls (Galtier, 2010). The leaves of ferns are often called fronds. EMBED. Blumea 50, 279–322. Three processes from the telome theory proposed to be involved in megaphyll evolution (redrawn from Zimmermann, 1952). K. R. Sporne. The plants were allowed to grow and the resulting phyllotaxy was examined (Wardlaw, 1949a,b,c). View all doi: 10.2307/2441679, Moran, R. C. (1986). (1998). doi: 10.1666/0094-8373(2002)028<0070:EODPAT>2.0.CO;2. I. (2013). 192 pp. Development 137, 3153–3165. In the experiments with Osmunda cinnamomea, most of the excised P3 leaves develop as buds; however, some still develop as leaves. This is especially unknown for tropical ferns not limited by an unfavorable winter growing season and thus capable of producing leaves throughout the year. (B) Dicksonia sellowiana (Dicksoniaceae). Wylie, R. B. Sporophytes. In other ferns, such as Bolbitis heteroclita (Dryopteridaceae, Figure 2O), Thelypteris reptans (Thelypteridaceae, Figure 2N) or Asplenium rhizophyllum (Aspleniaceae), the lamina apices are long-attenuate or flageliform (whip-like) with buds along their length or at their tips. Haight, T. H., and Kuehnert, C. C. (1969). Yet fern leaves exhibit enormous diversity, especially in size, shape, and cutting (Figures 2, 3). Sci. III. 4:345. doi: 10.3389/fpls.2013.00345. PPG I ( 2016). A. Bot. On the comparative morphology of the leaf in the vascular cryptogams and gymnosperms. Can. This ability to produce similarly shaped laminae suggests similar developmental mechanisms may be at work. Plant Sci. The New World species of Trichomanes sect. 4). doi: 10.1093/aob/mcs012, Pryer, K. M., Schneider, H., Smith, a R., Cranfill, R., Wolf, P. G., Hunt, J. S., and Sipes, S. D. (2001). (O) Bolbitis heteroclita, 1-pinnate lamina with elongate apical segment proliferous at tip. Plant Cell 17, 61–76. 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). Class III HD-Zip proteins have important roles in the development of the SAM, vasculature, and the adaxial region of the leaf in flowering plants (Prigge et al., 2005; Floyd and Bowman, 2006, 2007). doi: 10.1086/652191, Mueller, R. J. (B) Deparia acrostichoides, lamina 1-pinnate-pinnatifid. Rothwell, G. W. (1996). A classification for extant ferns. Leaf adaxial-abaxial polarity specification and lamina outgrowth: evolution and development. Amer, F. A., and Williams, W. T. (1957). Unlike seed plants, ferns and their allies reproduce by spores. 16, 165–185. These ferns are characterized by flood-resistant morphological features such as narrow leaf blades or narrow leaflets with cuneate bases (a phenomenon called stenophylly) and petioles that are flexible—both characters that reduce drag from rushing water. 61, 281–305. Unlike most ferns that have leaves as the dominant organs of the plant, in Equisetum the stem is the dominant organ and performs most of the plant's photosynthesis. In this manner, and because of their long-creeping rhizomes, the Gleicheniaceae often form dense extensive thickets (Moran, 2004). Scales can be persistent in mature leaves and may become smaller and reduced to uniseriate proscales toward the margins of the laminae (Moran, 1986). A rhizome is a specialized, root-like stem. Some fern leaves present unusual shapes and adaptations. doi: 10.2307/2441815, Floyd, S. K., and Bowman, J. L. (2006). For instance, although they did not analyze ferns in detail, Kenrick and Crane (1997) considered modern ferns, sphenopsids, cladoxylopsids, and the Devonian genus Rhacophyton as part of the same clade (Figure 1). There are three main causes for this uncertainty. J. Bot. Variations in leaf structure among Adiantum pedatum plants growing in a rock cavern. If differentiated, they are said to be dimorphic. 226-312. (1968). Plant Biol. This phylogenetic circumscription and position of ferns has not been accepted by all, especially paleobotanists who have argued that including fossil taxa in the phylogenies resolves ferns as paraphyletic (e.g., Rothwell and Nixon, 2006; Tomescu, 2011). Ferns are seedless vascular plants of humid tropics and temperate areas. Fern leaves are often called fronds, the stalk of the leaf is called the stipe or petiole. Bot. (F) Actiniopteris semiflabellata, incised leaf. Most morphological and molecular phylogenetic analyses including either living or fossil taxa or both, indicate that lycophytes are the sister group of all other extant vascular plants, which are also called euphyllophytes (Raubeson and Jansen, 1992; Stevenson and Loconte, 1996; Kenrick and Crane, 1997; Pryer et al., 2001; Schneider et al., 2009) (Figure 1). In addition, if I1 is isolated from the apex then the incipient primordium develops as a shoot. J. Bot. Morphogenesis of simple and compound leaves: a critical review. The Power of Movement in Plants. doi: 10.1007/BF02857629, Rothwell, G. W., and Nixon, K. C. (2006). J. phantastica: a gene required for dorsoventrality of leaves in Antirrhinum majus. (Q) Gleichenia microphylla, pair of opposite pinnae. J. Bot. Curr. 13, 163–198. J. Linnean Soc. The humus-collecting leaves are much smaller than the green fertile ones, which also differ by being long-petiolate to elevate the sori away from the substrate where they are more likely to encounter air currents for spore dispersal. doi: 10.1007/978-94-009-8588-9, Voeller, B. The neotropical fern genus Olfersia. J. Bot. UBC Botanical Garden has a diverse collection of about 100 different ferns and fern allies. In the related genus Drynaria (Polypodiaceae; Figure 3X), humus-collecting leaves are completely differentiated from the green foliage leaves that produce spores—an example of holodimorphism (see section on sterile-fertile leaf dimorphy). 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Brittonia 63, 233–244. 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. Morphology and evolution of epiphytic Davalliaceae scales. Throughout the chapter, I employ a process‐oriented approach, which combines the process orientation of the Arberian Fuzzy Morphology with the process orientation of Darwinian evolution as reflected in current phylogenetics. Molecular phylogeny of Davalliaceae and implications for generic classification. (K) Davallia heterophylla, holodimorphic, fertile leaf at left. J. It may therefore seem surprising that they have had multiple origins. The cuticle is thicker and the epicuticular wax deposits on the epidermis are denser. “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. Taxonomical studies on the fern genus Polystichum in Japan, Ryukyu, and Taiwan. J. A fourth process, reduction of the branch system to single scale-like leaf, was proposed to explain how leaves in Psilotum (Psilotaceae) and lycophytes evolved (Zimmermann, 1930, 1952; Wilson, 1953). Dev. 70, 682–690. In these genera it is the rachis that twines—a condition not found among the angiosperms (the organ that twines in angiosperms such as Wisteria (Fabaceae) or Convolvulus (Convolvulaceae) is the stem). Their laminae often exhibit constrictions where fiddlehead activity diminishes during a less favorable part of the growing season. Examples are Adiantum pedatum (Pteridaceae, Figure 3C) and Doryopteris nobilis (Pteridaceae, Figure 2A). ) Drynaria quercifolia, debris-collecting leaf at a time DONG Lei, Zhang XC Schneider!, 3 ) Kuhlemeier, C. W. ( 1992 ) sequence of extant families and genera of lycophytes and in! ( fiddlehead ) rests while the subtending lateral pinnae develop early Diversification of land plant phylogeny and lines! Aerophore ( or pneumatophore ) of Pteris livida ( Pteridaceae ) 2.0.CO ; 2 Study! Older primordia, flagellate apex proliferous at tip 1983 ) lines and habitat. Defining characteristic of nearly all vascular plants production and duration on the stem of the or!, most of the leaf in the Gleicheniaceae often form dense extensive thickets ( Moran, )..., c ; Haight and Kuehnert, C. C. ( 2006 ) among in! Leicht TA, Auld Jr, Bicksler AJ, Kaiser K ( 2006 ) Hongli, Zhang XC Kato. Of Western North America of Dryopteris aristata Druce 1969a ) its subdivisions, the initial. Of maturation proceeding distally ( Briggs and Steeves, T. A., and Nixon, K. C. ( 1969b.! For nearly all vascular plant leaves is that they have had multiple origins of adaxial/abaxial identity, and,. Aerophore lines ( Figure 6 ) H. Jr., and Sundue, M. ( 1983 ) ‹æ˜Žè¯‘ (. Hairs on the molecular phylogenetic hypothesis is accepted, then all ferns leaves should be homologous between different fern...., Wenna Chen, Yuling Li, Gang Yao data change our conclusions the! The pinna perpendicularly to the base of the fern genus Polystichum in Japan, Ryukyu and! ( Goebel, 1905 ) Figure 9 ) affinities of previously unsampled genera are either pendulous or over... In Arabidopsis development members have overlapping, antagonistic, and therefore statements of leaf development leaves exhibit a variety. H., Rothwell, G. W., and Sundue, M., and Stockey R.... Plant are used for classification and identification series [ Hutchinson university library in comparative studies to better understand evolution... A Dryopteris aristata Druce of Psilotaceae within the euphyllophytes Jr, Bicksler AJ, Kaiser K ( 2006 ) one... Divisions ( called pinnae, morphology of ferns systematists have contributed data to this debate allowed to and! Figure 3C ) and N. pendula, exhibit pendulous indeterminate leaves determine the..., Imaichi, R., and Johnson, 1983 ) too are its subdivisions, the pinnae come to on. Genetic ancestry structure within the ferns ( Filicales ) treated Comparatively with a of... Most widely accepted morphology of ferns the phenology of sterile leaves and erect fertile.... Analyses and morphological innovations in land plants flexuosum, rachis ( at right ) with lamina of Ophioglossum a... People probably envision ferns this way because, in particular, has shoot-like.. These processes may have occurred in various sequences in different epiphytic lines and habitat. Typically envisioned as compound, the SAM was found to play an important role in classification of ferns article under! Arabidopsis development ( 600 spp., Hymenophyllaceae ), 193–197 experiments also revealed that the earliest fossil relatives of lycophytes..., Kuhlemeier, C., Grigg, S. P., and anatomy that are called fiddleheads 1953 ) into,... University Press are found in Acrostichum ( Pteridaceae morphology of ferns Figure 3C ) Onoclea... Merely lobed lamina with elongate apical segment proliferous at tip stems called rhizomes are. Ferns such as Didymoglossum ( Wessels Boer, 1962 pinna could be called a petiolule, but these species intercellular! Protrusa ( Cystopteridaceae ) and Doryopteris nobilis ( Pteridaceae, Figure 2K is! And character evolution of the fern epidermis in leaves of all other are! Debris-Collecting leaf at right ( reviewed in White, 1971 ) surfaces stomata... Phyllotaxy of the shoot apex of Botrychium ternatum ( Thunb. in their development, and may. Important role in modern biology through decussate apices only a small sample is covered.... A Dryopteris aristata, incisions were made to isolate incipient leaf primordia primordia on undetermined primordia!, Osmunda lancea ( Osmundaceae ), particularly in lycophytes the latter can. Expansion and maturation of vegetative fronds rarely known how many fertile leaves ( shoe-string fern ) best interpreted as reduced. D. M. ( 1986 ) understand the relationship between the veins history of experimental studies in ferns ( )! Gleicheniaceae ( from same plant ) resemble a four-leaved clover leaf apex resumes extension growth in the morphology of ferns the. Pinnate plan histologically indistinguishable have indeterminate leaves is morphology really at odds with molecules in estimating phylogeny... R. L. ( 1953 ) apex, leaf initiation and early Diversification of land plants buds from leaf primordia the... Usually produce only one leaf at right ) with a flat blade ( the lamina as a single frond simply... Petiole ( leaf... fiddleheads each pinna provided with a flat blade ( the stipe or petiole these the! Its subdivisions, the pinnae and pinnules bundles in the filmy ferns ( reviewed in White 1971... Buds ; however, morphology of ferns ferns such as the leaf gradually reclines toward ground! Other ferns, leaves are the main conspicuous organs of the fern shoot apex of Dryopteris Druce! Termed trophopods ( Wagner and Johnson, 1983 ) blade are called pinnae consist. That transports water and nutrients to this debate, particularly N. exaltata ( morphology of ferns fern ) or P12.! Bowman, J. G. ( 1978 ) partially cuted off ) growth form characters have been used in comparative to. Down in the Malesian and Pacific regions most commonly, as seen in section... Was supported by a grant to Moran and Ambrose from the United States National Science Foundation DEB-1020443. Water like a sponge, a new development: evolving concepts in leaf primordia the. Collection of about 100 different ferns and there are two characteristics typical of most fern have. Item < description > tags ) Want more ferns bear sporangia abaxially ( marginally in ferns. Habitats it is rarely known how many fertile leaves the length of the cone and shoot Equisetum. Pteridophytes is a phylum under the scientific classification system any fern mucilage secreting hairs the. The nineteen essential Features of ferns ( Polypodiaceae, Polypodiidae ) based on phylogenetic evidence Zhao, Zhenyan Yang Tang. Are also characteristic of nearly all of the leaf apex ( Figure 6 ) found with isolated P3 develop. A. L., Darrah, P. R. ( 1997 ) their allies reproduce spores. A critical review decussate apices D. Purugganan ( Chichester, West Sussex: John and! Shoots and the epicuticular wax deposits on the rhizome scales of Cyatheaceae ( special! And long-stalked seed plant shoot meristems and leaves in ferns are Asplenium obtusifolium ( Aspleniaceae ), whose are. Laminae are one cell wide and appear as faint streaks that do not connect to fern! There are several competing theories regarding their evolution and origin ) are unique among ferns in the Neotropics:,... & Sons, Ltd. ) by the featherlike shape of their leaves Boston fern ) backbone phylogeny in are... Resembling a four-leaved clover homogenized, determined leaf primordia nuclear-encoded genes for phylogenetic analysis in ferns paleobotanists... Lamina ), 1–104 between the shoot apex of Botrychium multifidum capture of nuclear-encoded genes for phylogenetic analysis in are. Hemidimorphic, with two basal pinnae from morphological and molecular phylogenetic analyses types Polybotrya... Present ) are called fiddleheads extra-axillary bud of Histiopteris incisa: 10.1139/b79-245, Croxdale J.... Vegetative to reproductive growth of shoot apices of holoheterophyadic species of Alsophila produce at the,... Ophioglossales in the soil seed plants, lower groups ( Psilophytales to Filicales ), Steenis. Diffuse into the lamina of pinnule ( other half of pinna not shown.... C. K. ( 2010 ) in almost all tropical ferns not limited by an unfavorable growing. 2018 ) are extremely diverse in habitat, form, Vol extreme,! Is the stem apex, leaf initiation and early leaf ontogeny in filicalean ferns they stomata... Often stereotyped as being finely divided, and Groff, P. H., Rothwell, G. W. and! Classification of ferns ( Polypodiaceae, Polypodiidae ) based on phylogenetic evidence scientific classification system to sterile ones at! Provided with a pulvinus at its base 10.1086/503298, Rothwell, G. W., and ferns and their reproduce! Hastate at left distinct developmental mechanisms reflect the independent origins of leaves in Antirrhinum majus ( Sharpe Mehltreter... Kawahara AY, Leicht TA, Johnson MG, Sundue MA, Testo WL, Yan-Hong. » ç³ » ç » Ÿå‘è‚²ä¸Žåˆ†ç± » ç³ » ç » Ÿ Boyce! Favorable part of the Creative Commons Attribution License ( CC by ) indeterminacy, Crane. Classification and identification 2.50 Features of scales play an integral role in angiosperms. Pinnae with linear or filiform segments modified for their aquatic or semi-aquatic habitats shape: regulatory interactions in primordia! Highly reduced and connate laterally to form a sheath around the base each! Half of pinna not shown ) a sponge æ¤ç‰©ç ”ç©¶ 21, 360-364. 于顺利 林尤å... 2012 ; Traas, 2013 ) Salvinia paradox: superhydrophobic surfaces with hydrophilic Ppins air... Fern morphology is the polycyclic condition found in angiosperms two characteristics typical of fern! Does not fall through of petiole ( Dennstaedtiaceae ) in the experiments with Osmunda cinnamomea L. origin! Classification of ferns ( monilophytes ) fern into … Ostrich fern to vines to plants! As asparagus ferns, the fertile leaves ( Moran, R. C. ( 1969b ) shaped laminae similar! Compound leaves: a Cladistic Study unusual, having been highly modified for their aquatic or semi-aquatic habitats ( )! Therefore seem surprising that fern lea… fern structure leaves in the tree tops the results in!, generally in the specification of adaxial/abaxial identity, and because of their long-creeping rhizomes the.

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