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Glossary of Botanical Terms Used in this Volume Accepted terms are indicated by boldface. Within the definitions, italics (when not names of genera) indicate terms that are defined in this glossary. (Most definitions have been taken or adapted with permission from Lellinger, D. B. 2002. A modern multilingual glossary for taxonomic pteridology. Pteridologia 3. [Washington, D.C.]: American Society, Inc.) acroscopic - facing or directed toward the apex of the axis Woodwardia species; synonyms , bulblet; see also and on which the structure is borne; cf. basiscopic. proliferous. acrostichoid - of sori, producing sporangia apparently or bulblet - see bulbil actually spread across the surface of the fertile lamina, - the part of a that contains the . usually densely so. catadromous - with the basiscopic pinnule and/or vein group of actinostele (adjective actinostelic) - a radially arranged, the pinna the first to depart from its axis; cf. anadromous. lobed protostele (as seen in cross section) associated with microphylls, commonly found in the . clypeate - shaped like a rounded shield. aerophore - a spot, swelling, or fingerlike projection of coenosorus (plural coenosori) - a compound composed of thin-walled cells found along the or in the pinna or several contiguous sori fused end-to-end. pinnule axils of some , especially ; commissure (adjective commissural) - place where a more prominent on young . commissural vein joins parallel, otherwise free veins along the alate - of spores, with the outer wall (exospore or perispore) lamina margin and often underlies a continuous, marginal raised in a pattern of narrow, winglike ridges surrounding coenosorus. depressions. costa (plural costae, adjective costal, costate) - the major axis of alete - of spores, spherical, lacking a laesura, and containing a pinna; synonyms midrib, midvein. more sets of chromosomes than normal spores, found costate - of scales, with a central line of cells different (usually regularly or occasionally in some apomictic ferns. darker) from the more marginal cells, as in certain anadromous - with the basal pinnule and/or vein group of and ; of spores, with the outer wall (exospore or the pinna directed toward the apex; cf. catadromous. perispore) raised in a pattern of relatively broad ridges with rounded crests. anisophyllous - bearing fronds or microphylls of unequal size, but of a single shape. costule (adjective costular, costulate) - the major axis of a pinnule; synonyms midrib, midvein, rachilla. annulus (plural annuli, adjective annular) - a row or patch of partially or entirely thick-walled cells of the capsule of the crozier - see fiddlehead leptosporangium which contracts and forces the capsule to ctenitoid - of hairs, with adjacent cells collapsed at right angles open and to discharge its spores. to each other and often the end walls of the cells thickened or (adjective antheridial) - the male organ dark-colored, as on some fronds of Ctenitis. of pteridophytes borne on the and producing dictyostele (adjective dictyostelic) - a siphonostele with more spermatozoids. than one parenchymatic gap at a single level (as seen in cross (adjective apomictic) - the formation of a section). from a gametophyte by direct, asexual echinate - of spores, with the outer wall (exospore or perispore) development, rather than by fertilization of . raised in a pattern of long spines. apophysis (plural apophyses) - a swelling on which a sorus epispore - a particular, outer, layer of the is located. wall that is external to, but partially attached to, the exospore in (plural archegonia) - the female some heterosporous ferns and . of pteridophytes borne on the gametophyte and producing exine - see exospore . exospore - the principal sporopollenin layer of the spore wall areole (adjective areolate) - an area surrounded by internal to the perispore; synonym exine. anastomosing veins. false indusium - an introrse, reflexed or revolute, often baculate - of spores, bearing cylindrical projections more modified lamina margin that protects young sporangia. than 1 μm long and less in diam., and usually with an obtuse false vein - an elongate series of thickened cells appearing to be to rounded apex. a vein, but not connected to true veins and not functioning as a basiscopic - facing or directed toward the base of the axis vein; found in the laminae of certain Aspleniaceae, on which the structure is borne; cf. acroscopic. , , , and blade - see lamina Selaginellaceae. bud - see bulbil ferns - the pteridophytes excluding the Isoëtaceae, Lycopodiaceae, and Selaginellaceae. bulbil (adjective bulbiliferous) - a small, usually persistent, ± globose, usually hairy or scaly, asexual borne on fiddlehead - the young, unexpanded, circinate apex of a fern a , , or frond and capable or not capable of frond; synonym crozier. forming a plantlet, as in, e.g., certain Asplenium, Bolbitis, frond - the photosynthetic organ of ferns, usually consisting of a , , , Polystichum, Tectaria, and stipe and lamina; synonyms , megaphyll.

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gametophyte - an inconspicuous, non-vascular stage in the lumen (plural lumina) - the central cavity of a cell, especially cycle of a that bears gametangia with applied to the cells of clathrate rhizome scales. gametes. In homosporous pteridophytes, they are either - collectively the Isoëtaceae, Lycopodiaceae, and surficial, thin, chlorophyllous, and various in shape Selaginellaceae. (filamentous, ribbonlike, heart-shaped, or somewhat stellate) massula (plural massulae) - in , a structure derived from or subterranean, massive, achlorophyllous, and globose, the contents of the microsporocarp that contains the cylindrical, or branched. In heterosporous pteridophytes, and has glochidia (minute barbed hairs) protruding from its they are much reduced structures borne (and partially surface. developing) within spore walls; synonyms prothallium, . megagametophyte - in heterosporous pteridophytes, a female gametophyte borne within a megasporangium and bearing one or (plural gemmae, adjective gemmiferous) - a gemma more archegonia. structurally specialized, asexual propagule found on some that detaches and forms a new gametophyte; megaphyll - see frond also used in the sense of bulbil in some African or European megasporangium (plural megasporangia) - a sporangium fern literature. bearing . goniopteroid - of veins, a system of excurrent veinlets - a large spore of the heterosporous pteridophytes connected to more distal vein unions, or to a translucent line Azolla, Isoëtes, , , Regnellidium, , and leading to a sinus, thus forming oblique rhomboid areoles. that produces a female gametophyte. hemitelioid - shaped like a shallow saucer or fan, usually megasporocarp - a that bears megasporangia. firm and fully or partially surrounding the sorus, typical of megasporophyll - a fertile microphyll bearing or subtending a certain . megasporangium. heterosporous (antonym homosporous) - producing microgametophyte - in heterosporous pteridophytes, a male spores of two sizes, each of which develops gametophytes gametophyte borne within a microsporangium and bearing one or having gametangia of a single sex. more antheridia. hydathode - a dark or sometimes pale area of the microphyll - the photosynthetic organ of the lycophytes, on the adaxial surface of the lamina that coincides with the , and , always lacking a stipe, often small endings of the veins and exudes water, salts, etc. and generally supplied with a single ; usually indusium (plural indusia, adjective indusiate) - a usually associated with a protostele or siphonostele. thin, often scalelike, epidermal membrane subtending and/or microsporangium (plural ) - in heterosporous covering the sorus, that partially or fully protects the young pteridophytes, a sporangium bearing microspores. sporangia. microsporocarp - a sporocarp that bears microsporangia. intrastelar canal - a channel occurring within a stele. microsporophyll - a fertile microphyll bearing or subtending a isodromous - with the basal pinnules and/or vein groups of microsporangium. the pinnae strictly opposite. midrib, midvein - see costa, costule, and rachis isophyllous - having fronds or microphylls of a single size monolete - of spores, bilaterally symmetric with a linear, and shape. unbranched laesura. laesura (plural laesurae) - the simple, elongate or triradiate, murus (plural muri, adjective muriform) - of spores, an scar on the surface of pteridophyte spores; synonym suture. elongate, wall-like protuberance. lamina (plural laminae, adjective laminate) - the expanded paraphysis (plural paraphyses) - a minute, unicellular or portion of a frond, usually consisting of a rachis or costa, multicellular (resembling a simple hair), usually elongate and other axes or lateral veins, and expanded lamina ; sometimes glandular structure borne on the soral , on synonym blade. the sporangium capsule or . leaf - see frond perine - see perispore leptosporangium (adjective leptosporangiate) - a perispore - the outermost, sporopollenin layer of the spore wall thin-walled, thin-pedicelled sporangium bearing usually 64 that is deposited on the exospore; synonym perine. spores (32 in apomixises of the Dryopteridaceae, 128-512 in phyllopodium (plural phyllopodia) - in ferns with articulate the , and 256 in the Schizaeaceae) and formed stipes, that portion of the stipe proximal to the articulation that usually from a single epidermal initial cell. remains attached to the rhizome. Phyllopodia are especially ligule (adjective ligulate) - a small, tonguelike, often prominent and stipelike in, e.g., Elaphoglossum and Oleandra but triangular appendage located near the microphyll base (just low and more like the rhizome and knoblike in , distal to the sporangium on the adaxial surface of the Davalliaceae, and many Polypodiaceae. microsporophyll); it is persistent in Isoëtes. pinna (plural pinnae, adjective pinnate) - a stalked or sessile, lophate - of spores, with the outer wall (exospore or primary division of a compound lamina that is at least narrowed at perispore) raised in a pattern of ridges (lophae) surrounding the base. depressions. pinnule - a stalked or sessile division of a pinna that is at least narrowed at the base.

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plectostele (adjective plectostelic) - a vascular cylinder sporangiophore - a greatly transformed, peltate with the appearing to be ± parallel plates (as bearing a ring of ca. 6 sporangia facing the axis of the of seen in cross section), associated with microphylls, found in Equisetum. many Lycopodiaceae. sporangium (plural sporangia) - the spore-producing structure primordium - a part (e.g., a frond) in its most rudimentary of pteridophytes. form or stage of development. spore - a spherical, tetrahedral, or reniform, often elaborately proliferous - forming bulbils or plantlets, often on parts ornamented, reproductive cell that is produced within the that normally have another function such as (e.g., sporangium and germinates to form a gametophyte. ), stems and branchlets (e.g., Huperzia), rachises sporocarp - in Marsilea, the hard, short- to long-pedunculate, (e.g., most Bolbitis, Diplazium, Dryopteris, some Hymenophyllaceae, Polystichum, Tectaria, and nutlike structure containing the sporangia, apparently a highly Woodwardia) or lamina margin (e.g., some Asplenium). modified leaflet; in Azolla and Salvinia, a thin, short-stalked, globose structure containing the sporangia, apparently a modified prothallium, prothallus (plural prothallia, prothalli) - see indusium. gametophyte sporophore - the fertile portions of a hemidimorphic frond as in protostele (adjective protostelic) - a simple vascular the . cylinder that lacks a pith of parenchyma in the center and is without parenchymatic gaps. sporophyll - in ferns, a frond bearing sporangia; in the lycophytes, Equisetaceae, and Psilotaceae, a microphyll pteridophytes - non--bearing vascular ; ferns and subtending a sporangium; in Isoëtes, an elongate microphyll lycophytes collectively. bearing a sporangium within its base. pulvinus (plural pulvini) - a swollen structure at the base of (adjective stelic) - the vascular and associated tissues of a a frond or at the base of pinnae, particularly common in the stele Marattiaceae. rhizome or other type of stem; see also dictyostele, protostele, siphonostele, and solenostele. rachilla - see costule stipe (adjective stipitate) - the structure of a frond that connects rachis - the principal, central axis of a pinnatifid or more the base of the lamina to the point of its attachment to the compound lamina. rhizome. receptacle - the point or region of the lamina tissue, often stipicel - a term sometimes used for the stalk of a pinna or thickened and amply supplied by one or more veins, that pinnule. produces sporangia and sometimes paraphyses and/or sporangiasters. stipule - in the Marattiaceae, each one of a pair of lateral, fleshy, -bearing, persistent, partially or entirely vascularized - an elongate, non-vascularized, uni- or outgrowths of the rhizome that clasp the base of the stipe and that paucicellular structure that serves to anchor the gametophyte and to absorb water and nutrients from the substrate. are capable of vegetatively reproducing the plants; in the Ophioglossaceae, merely the remnants of the older stipe base that rhizome - in pteridophytes, a scaly or hairy (rarely originally enclosed and protected the younger, less developed glandular or glabrous) anchoring stem that bears roots and fronds. fronds. strobilus (plural strobili) - in the lycophytes and Equisetaceae, a rhizophore - a specialized, aerial root of Selaginella that compact reproductive structure borne at the tips of branches or arises in the axils of stems and branches repeatedly when in axes consisting of a central axis bearing closely spaced, spirally contact with the substrate; it may occasionally differentiate arranged or sporangiophores. into a stem, rather than a root. suture - see laesura. rugate - of spores, bearing muri that are wide, rounded, and non-anastomosing and that do not form areoles. synangium (plural synangia) - a group of sporangia partially or sinus membrane - an elongate translucent membrane entirely fused laterally, as in and Marattiaceae. below the sinus in a pinnule. trilete - of spores, radially symmetric (spherical or tetrahedral) siphonostele (adjective siphonostelic) - a vascular cylinder with a laesura with three radiating branches. that has a pith of parenchyma in the center and on trophophyll - a vegetative, nutrient-producing frond or both the outside and inside of the cylinder, or (in the microphyll. Osmundaceae) only on the outside of the cylinder. trophopod - the enlarged, persistent, basal portion of a stipe that solenostele (adjective solenostelic) - a siphonostele with functions as a . only one parenchymatic gap at a single level (as seen in cross valve - an involucral lobe, especially in Hymenophyllum and section). some Dicksoniaceae and Dennstaedtiaceae; also, each half of a soriferous - bearing sori. sporangium that is divided into halves, as in the Lycopodiaceae sorophore - the gelatinous, sporangium-bearing ring and Osmundaceae. produced by sporocarp (as in the ) during vascular bundle - an elongate strand of conducting cells ( . tracheids and phloem sieve cells) that serve to conduct water, sorus (plural sori, adjective soral) - a cluster of sporangia. mineral nutrients, and photosynthetic products. spermatozoid - a motile male sex cell () produced velum - in Isoëtes, the membrane covering part or all of the in the antheridium. sporangium-containing cavity (fovea) in the base of a microphyll.