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81 Vascular Plant Diversity f 80 CHAPTER 4 EVOLUTION AND DIVERSITY OF VASCULAR PLANTS UNIT II EVOLUTION AND DIVERSITY OF PLANTS 81 LYCOPODIOPHYTA Gleicheniales Polypodiales LYCOPODIOPSIDA Dipteridaceae (2/Il) Aspleniaceae (1—10/700+) Lycopodiaceae (5/300) Gleicheniaceae (6/125) Blechnaceae (9/200) ISOETOPSIDA Matoniaceae (2/4) Davalliaceae (4—5/65) Isoetaceae (1/200) Schizaeales Dennstaedtiaceae (11/170) Selaginellaceae (1/700) Anemiaceae (1/100+) Dryopteridaceae (40—45/1700) EUPHYLLOPHYTA Lygodiaceae (1/25) Lindsaeaceae (8/200) MONILOPHYTA Schizaeaceae (2/30) Lomariopsidaceae (4/70) EQifiSETOPSIDA Salviniales Oleandraceae (1/40) Equisetaceae (1/15) Marsileaceae (3/75) Onocleaceae (4/5) PSILOTOPSIDA Salviniaceae (2/16) Polypodiaceae (56/1200) Ophioglossaceae (4/55—80) Cyatheales Pteridaceae (50/950) Psilotaceae (2/17) Cibotiaceae (1/11) Saccolomataceae (1/12) MARATTIOPSIDA Culcitaceae (1/2) Tectariaceae (3—15/230) Marattiaceae (6/80) Cyatheaceae (4/600+) Thelypteridaceae (5—30/950) POLYPODIOPSIDA Dicksoniaceae (3/30) Woodsiaceae (15/700) Osmundales Loxomataceae (2/2) central vascular cylinder Osmundaceae (3/20) Metaxyaceae (1/2) SPERMATOPHYTA (See Chapter 5) Hymenophyllales Plagiogyriaceae (1/15) FIGURE 4.9 Anatomy of the root, an apomorphy of the vascular plants. A. Root whole mount. B. Root longitudinal-section. C. Whole Hymenophyllaceae (9/600) Thyrsopteridaceae (1/1) root cross-section. D. Close-up of central vascular cylinder, showing tissues. TABLE 4.1 Taxonomic groups of Tracheophyta, vascular plants (minus those of Spermatophyta, seed plants). Classes, orders, and family names after Smith et al. (2006). Higher groups (traditionally treated as phyla) after Cantino et al. (2007). Families in bold are described in found today in the Selaginellaceae of the lycophytes and all the pericycle or endodermis. Lateral roots penetrate the tis detail. Number of genera and species (often approximate), respectively, are indicated in parentheses, separated by slash mark. monilophytes (discussed later). In the Lycopodiaceae, sues of the cortex before exiting to the outside. Isoetaceae, and seed plants (see Chapter 5), the apical mer Numerous modifications of roots have evolved, most of laesura (Figure 4.1 istem is complex, consisting of a group of continuously divid these restricted to the flowering plants (see Chapter 9). Roots VASCULAR PLANT DIVERSITY spores, with a 3-branched lA); 2) monolete ing cells. of many, if not most, vascular plants have an interesting sym spores, with a laesura that is linear and unbranched (Figure Roots are characterized by several anatomical features. biotic interaction with various species of fungi; this associa A classification scheme of vascular plants, after Smith et al. 4.1 lB); and 3) alete, lacking any evidence of a laesura. First, the apical meristem is covered on the outside by a tion between the two is known as mycorrhizae. The fungal (2006) and Cantino et al. (2007), is seen in Table 4.1. Of the rootcap (also called a calyptra; Figure 4.9A,B); stems lack component of mycorrhizae appears to aid the plant in both tremendous diversity of vascular plants that have arisen since RHYN IOPHYTES such a cell layer. The rootcap functions both to protect the increasing overall surface area for water and mineral absorp their first appearance some 400 million years ago, only the Rhyniophytes are a paraphyletic assemblage that included root apical meristem from mechanical damage as the root tion and increasing the efficiency of selective mineral absorp major lineages will be described here. These include the the first land plants with branched sporophytic axes, some of grows into the soil and to provide lubrication as the outer cells tion, such as of phosphorus. The fungus benefits in obtaining rhyniophytes, known only from fossils, plus clades that have which (but not all) also had vascular tissue. Rhyniophytes slough off. Second, with the exception of the Psilotopsida photosynthates (sugars and other nutrients) from the plant. modern-day descendants: the Lycopodiophyta (lycophytes) incLude the genus Rhynia (Figure 4.12A,B), a well-known (Psilotales and Ophioglossales), the epidermal cells away from and Euphyllophyta (euphyllophytes; Figure 4.1, Table 4.1). vascular plant from the early Devonian, ca. 4 16—369 million the root tip develop hairlike extensions called root hairs See Bierhorst (1971) and Foster and Gifford (1974) for gen years ago. Rhyniophyte sporophytes consisted of dichoto (Figure 4.9A); these are absent from stems (although under eral information on vascular plant morphology. mously branching axes bearing terminal sporangia that ground stems of the Psilotales bear rhizoids, which resemble Features that have been used to classify vascular plants dehisced longitudinally. root hairs). Root hairs function to greatly increase the surface include sporophyte vegetative morphology (branching pat area available for water and mineral absorption. Third, roots tern, leaf type/shape/arrangement/venation, stem and leaf always have a central vascular cylinder (Figure 4.9C,D). As anatomy), life cycle and reproductive morphology (homo in stems, the mostly parenchymatous region between the vas spory/heterospory, sporophyll morphology, sporangium culature and epidermis is called the cortex (Figure 4.9C); the shape/dehiscence/attachment, spore morphology), and game center of the vascular cylinder, if vascular tissue is lacking, is tophyte morphology (whether green and photosynthetic or called a pith. Fourth, the vascular cylinder of roots is sur nongreen and saprophytic or mycorrhizal). Spore morphol rounded by an endodermis with Casparian strips (Figure ogy in particular has been useful in the classification of vas 4.9D). As with some stems, the endodermis in roots selec cular plant groups. (See Chapter 12.) Features include spore tively controls which chemicals are and are not absorbed by size, shape (e.g., reniform, tetrahedral, globose), sculpturing the plant, functioning in selective absorption. (An undifferen patterns, and whether green (photosynthetic) or not. One major tiated layer internal to the endodermis, called the pericycle, spore feature is related to the laesura (plural laesurae), the is also typically present.) Fifth, roots generally have endog differentially thickened wall region corresponding to the tetrad FIGURE 4.11 MONILOPHYTA. Spore morphology. A. Spore immature following scar triangularis, Pteridaceae). B. Spore enous lateral roots (Figure 4.10), in which new lateral roots FIGURE 4.10 Root cross-section (Lilium sp.), showing endoge attachment scar on each of the four spores with trilete (Penragramnw Aspleniaceae). originate by means of actively growing meristems, arising at nous lateral root, a charactenstic of vascular plant roots. meiosis. Three basic spore types are recognized: 1) trilete with monolete scar (Asplenium nidus, r 82 CHAPTER 4 EVOLUTION AND DIVERSITY OF VASCULAR PLANTS UNIT II EVOLUTION AND DIVERSITY OF PLANTS 83 Rhynia stem axes leaf cortex (lycophyll) protoxylem scars (exarch) ‘cc B ‘I I lycophyll leaf (lycophyll) stem vasculature (no leaf gap) B phloem AL C A vascular tissue FIGURE 4.12 A—B. Rhyniophytes. A. FIGURE 4.14 A,B. Lycopodium stem cross-section Reconstruction of Rhynia major, an early, exinct vascular plant. Note erect, showing protoxylem that is exarch (to periphery of stem). C. Lycophyll structure. leaves) branched stem (without bearing terminal sporangia. IReproduced from Kidston, and R. W. H. Lang. 1921. Transactions of the Royal Society of Edinburgh. vol. 52(4): 831—902.] B. Rhynia stem axes embedded in “Rhynie” chert. C—E. Lycophytes. C—D. Sigillaria, an extinct, woody lycophyte. C. Stem cross-section showing outer wood. D. Fossil metaxylem (i.e., away from the stem center; Figure 4.14A,B). Although all vascular plants impression of lycophyll leaf showing single vein. E. Fossil cast have shoots, fossil evidence an extinct, of Lepidodendron, woody, tree-sized lycophyte. Note lycophyll scars. Fourth, lycophytes, at least ancestrally, have sporangia that suggests that shoot systems evolved independently in the are dosiventral (i.e., flattened and having a dorsal, upper, and lycophytes and euphyllophytes (see later discussion), because ventral, lower, surface) Rhyniophytes ancestrally lacked both and dehisce transversely relative to their associated leaves evolved independently. Lycophylls roots and a leaf- roots usually have an endarch protoxylem. Protoxylem bearing shoot system; these the axis of the stem or subtending leaf (see Lycopodium, possibly originated from the transformation of small append two features evolved later, refers to the first tracheary cells that develop within a patch of Figure 4.1 5E). Fifth, lycophytes have sporophytic leaves, ages called enations prior to or within the lycophyte and euphyllophyte lineages (found, e.g., in fossil zosterophyllo xylem and that are typically smaller and have thinner cell (Figure 4.1). The usually just called “leaves.” (Although some liverworts and phytes and relatives), which are external, peg-like appendages stems of rhyniophytes were protostelic walls than the later formed metaxylem. In the roots of lyco all mosses have “leaves,” these occur on gametophytes only that lack vascular tissue. Lycophylls may have evolved (Figure 4.7) in which the first-formed xylem (known as pro via phytes, the protoxylem forms in a position interior to the toxylem) was “centrarch” (positioned and are not strictly homologous with the sporophytic leaves the development of vasculature tissue leading from the stem at the center). metaxylem (i.e.,
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