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BOT 201

Plant Systematics 2 Ed. Michael G. Simpson


CHAPTER 3 Viridiplantae (Chlorobionta) Chlorophytes Green Algae Streptophytes Charophytes Embryophytes Liverworts Mosses Bryophytes Hornworts Tracheophytes (vascular plants) Viridiplantate/Chlorobionta Monophyletic group Green algae and land plants (embryophytes) Cellulosic cell wall Specialized type of chloroplast * have chlorophyll a & b; thylakoids stacked into grana; manufactures true starch Cellulose Polysaccharide Glucose sugar units are bonded in the beta-1,4 position (1,4-glucopyranoside) Secreted outside the plasma membrane as microscopic fiber-like units (microfibrils) Imparts rigidity to the cell Thylakoids chlorophyll-containing membranes Grana stacks of thylakoids Starch polymer of glucose sugar units (polysaccharide); chemically bonded in the alpha-1,4 position Endosymbiosis intracellular cohabitation of an independently living, unicellular prokaryote inside a eukaryotic cell Viridiplantae: 1. Chlorophyceae green algae *green algae Paraphyletic primarily aquatic viridiplantae (chlorophytes + non-land plant streptophytes) 2. Streptophyceae Oogamy egg (nonflagellate, larger) and sperm cell; in all land plants Production of flagellate, haploid gametes isomorphic primitive type of green plane sexual reproduction Fertilization union of gametes Haplontic life cycle: zygote (free-living) meiosis 4 haploid spores germination haploid individual Charophytes w/in the streptophytes plasmodesmata Plasmodesmata pores in the primary (1) cell wall through w/c membranes traverse bet. cells transfer of compounds bet. cells, rapid transport of solutes Charales Closest living relatives to the land plants Some capable of precipitating CaCO3 as an outer layer Grow by means of a single apical shell Lack true parenchyma Specialized male and female gametangia (antheridia and oogonia) have an outer layer of sterile cells Oogonia spirally arranged group of outer tube cells Retain the egg and zygote on the plant body Embryophyta (land plants) Silurian period (400 million years ago) first colonization of plants on land Embryo immature Sporophyte that is attached to or surrounded by the gametophyte; can remain dormant for a
Chapter 3: Evolution & Diversity of Green and Land Plants Chapter 4: Evolution & Diversity of Vascular Plants
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period of time and will begin to grow after the proper environmental conditions are met Sporophyte separate diploid phase in the life cycle of all land plants; forming from the zygote by the delay of meiosis and spore production a. Sporangium spore-producing region; contains sporogenous tissue matures into sporocytes b. Sporangial wall envelopes the sporangium; 1/more layers of sterile cells c. Sporocytes cells that undergo meiotic division; each sporocyte produces 4 haploid spores Gametophyte haploid, gamete-producing part of the life cycle Haplodiplontic life cycle (alternation of generations) Sporophyte as a separate phase of the life cycle large increase in spore production Sporophytes diploid ploidy level: a. Potentially prevents the expression of recessive, deleterious alleles b. Permits increased genetic variability in the Sporophyte generation Cuticle protective layer that is secreted to the outside of the cells of the epidermis; prevention of desiccation Epidermis provides mechanical protection of inner tissue; inhibit water loss Cutin polymer of fatty acids; sealant prevents excess water loss; impregnates the outer cellulosic cell walls of epidermal cells Parenchyma tissue o Solid mass of tissue derived from the apical meristem region o Apical meristem region of actively dividing cells; 1 cell for liverworts, hornworts, mosses, monilophytes o Consists of cells that most resemble the unspecialized, undifferentiated cells of actively dividing meristematic tissue o Parenchyma cells: a. Elongate to isodiametric b. Primary (1) cell wall only c. Living at maturity and potentially capable of continued cell divisions o Metabolic activities o May further differentiate into other specialized cell types Middle lamella pectic-rich layer that develops bet. the primary cell walls of adjacent cells; binds adjacent cells together Antheridium specialized gametangium of the haploid gametophyte; contains the sperm-producing cells; surrounded antheridial wall (sterile) Sterile jacket layer protects the developing gametes from desiccation Archegonium specialized female gametangium; venter (outer layer of sterile cells) surrounds the egg o Egg cell located inside and at the base of the archegonium o Ventral canal cell above the egg o Neck canal cells above the ventral canal cell o Protect the developing egg; fertilization; site for embryo/Sporophyte devt; establishment of a nutritional dependence of the Sporophyte upon gametophytic tissue

Iigo Atienza 2Biology-3

BOT 201
Plant Systematics 2 Ed. Michael G. Simpson
Presence of various ultrastructural modifications of sperm cells Flavonoid chemical compounds Proliferation of heat shock proteins Bryophytes nonvascular land plants Liverworts, mosses, hornworts Paraphyletic Gametophyte dominant, photosynthetic, persistent, and free-living phase Sporophyte small, ephemeral, attached to and nutritionally dependent upon the gametophyte Liverworts (Hepaticae) Monophyletic Apomorphies: 1. Distinctive oil bodies 2. Elaters elongate, nonsporogenous cells w/ spiral wall thickenings; inside the sporangium; hygroscopic change shape/move in response to moisture content; spore dispersal Morphological types: 1. Thalloid thallus (flattened mass of tissue) o Rhizoids uniseriate, filamentous processes that function in anchorage and absorption o Pores upper surface of the thallus gas exchange; not true stomata no regulating guard cells o Some have symbiotic relationship w/ Cyanobacteria o Gemma cups upper surface; Marchantia; contain propagules (gemmae); asexual reproduction 2. Leafy o Gametophytes stem axis bearing 3 rows of thin leaves o Stem prostrate o Most liverworts leaves modified; upper 2 rows of leaves are larger, lowermost row are reduced o Some more erect; similar 3 rows of leaves Antheridia and archegonia develop on the gametophyte Antheridiophores stalked, peltate structure bearing antheridia Archegoniophores stalked, peltate structure bearing archegonia Fertilization zygote mitosis diploid embryo mature diploid Sporophyte Sporophyte small, nonphotosynthetic, short lived; consists mostly of sporangium/capsule Mosses (Musci) Most speciose and diverse Stomates specialized epidermal cells generally found on leaves, sometimes on stems; found in mosses, hornworts, vascular plants; regulation of gas exchange o 2 guard cells chloroplast-containing; changes in Turgor pressure, can increase/decrease the size of the opening (stoma) bet. them; one/more ridge-like deposits on the side facing the stoma rich in suberin o Suberin waxy, water-resistant substance; seal the stoma Aerial Sporophyte axis mosses, hornworts, vascular plants; precursor to the evolution of the dominant, aerial sporophytic stem in vascular plants Apomorphies: 1. Hydroids specialized conductive cells; water conduction 2. Leptoids sugar conduction 3. Perine layer thick outer layer of spores; prevent excess desiccation Trilete mark bryophytes; three-lined structure on the spore wall; scar of attachment of the adjacent 3 spores of the 4 spores produced at meiosis Gametophytes always leafy Leaves have costa composed of conductive cells; resemble a true vein Antheridia, archegonia produced at the apex of gametophytic stems Calyptra apical archegonial tissue; protects the young sporophyte apex Stipe stalk of the sporophyte Sporangium/capsule apex of the stipe; have specialized mechanism of dehiscence Operculum falls off the capsule apex time of spore release Peristome teeth hygroscopic; as the capsule dries up, the teeth retract release of spores Protonema filamentous structure; result of initial devt of gametophyte; ancestral vestige; after growth turns into parenchymatous gametophyte Sphagnum peat moss; grows in wet bogs; makes the surrounding water acidic; 2 cell types: chlorophyllous cells, hyaline cells Chlorophyllous cells form a network Hyaline cells large, clear; pores; helical thickenings Hornworts (Anthocerotae) Monophyletic Similar to thalloid liverworts in gametophyte morphology Lack pores Symbiotic relationship w/ Cyanobacteria; lives inside cavities of the thallus Sporophyte aerial, elongate, cylindrical, photosynthetic; indeterminate growth basal intercalary meristem Apomorphies: 1. Intercalary meristem region of actively dividing cells near the base of the sporophyte; surrounded by a protective collar of gametophytic tissue 2. Columella central column of sterile tissue 3. Pseudo-elaters groups of cohering, nonsporogenous, elongate, hygroscopic cells
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Chapter 3: Evolution & Diversity of Green and Land Plants Chapter 4: Evolution & Diversity of Vascular Plants

Iigo Atienza 2Biology-3

BOT 201
Plant Systematics 2 Ed. Michael G. Simpson
CHAPTER 4 Vascular plants monophyletic subgroup of the land plants Apomorphies: 1. Independent, long-lived Sporophyte 2. Branched Sporophyte 3. Lignified secondary walls w/ pits 4. Sclerenchyma structural support 5. Tracheary elements (xylem) water transport 6. Sieve elements (phloem) sugar transport 7. Endodermis selective transfer of compounds 8. Roots anchorage; absorption of water and nutrients Haplodiplontic alternation of generations haploid gametophyte; diploid Sporophyte Sporophyte dominant, free-living, photosynthetic, relatively persistent Gametophyte ancestrally free-living, may be photosynthetic, smaller, short-lived Land plants sporophyte is initially attached to and nutritionally dependent upon the gametophyte Vascular plants sporophyte soon grows larger and becomes nutritionally independent, w/ the subsequent death of the gametophyte Stems (sporophytic axes) branched; bear multiple sporangia Polysporangiomorpha fossil and extant taxa having branched sporophytic stems and multiple sporangia Pan-tracheophyta all descendents exclusive of the liverworts, mosses, and hornworts Dichotomous branching; apical meristem splits into two, equal meristems Pseudomonopodial modified growth pattern; starts out dichotomous; 1 branch becomes dominant, 1 branch appears lateral Sporophytic stems supportive organs; elevates reproductive organs and leaves; conductive organ Lignin complex polymer of phenolic compounds; incorporated into an additional cell wall layer (secondary *2+ wall) in some specialized cells Secondary walls secreted to the outside of the plasma membrane after the primary wall has been secreted; thicker; cellulosic; lignin is secreted into the space bet. cellulose microfibrils Pits holes in the secondary wall; occurs in pairs opposite the sites of plasmodesmata; allow chemical communication bet. cells; specialized function in xylem Primary pit field group of plasmodesmata Sclerenchyma Nonconductive cells Thick, lignified secondary cell wall w/ pit Dead at maturity 1. Fibers long, narrow cells w/ sharply tapering end walls; mechanical support; occur in groups/bundles; may occur independently of vascular tissue 2. Sclereids isodiametric to irregular/branched in shape; structural support; help deter herbivory in some plants Collenchyma live cells w/ unevenly thickened, pectic-rich, primary cell walls; not an apomorphy for vascular plants Tracheary elements (of xylem) Specialized cells that function in water and mineral conduction from the roots to other parts of the plant Elongate, dead at maturity, lignified 2 cell walls Joined end-to-end, forming a tube-like continuum Xylem tissue consisting of tracheary elements, w/ parenchyma and some sclerenchyma 1. Tracheids imperforate water and minerals flow bet. adjacent cells through the primary cell walls at pit-pairs; primitive type 2. Vessel members perforate 1/more continuous holes/perforations w/ no intervening 1 and 2 wall bet. adjacent cells through which water and minerals may pass *perforation plate contact area of 2 adjacent vessel members; compound (several perforations) or simple (1 opening) Sieve elements (of phloem) Conduction of sugars Phloem tissue consisting of sieve elements, w/ parenchyma and some sclerenchyma Elongate cells having only a 1 wall w/ no lignified 2 cell wall 1 wall has specialized spores w/c are aggregated together into sieve areas Each pore of the sieve area continuous hole in the 1 cell wall that is lined w/ callose Callose polysaccharide composed of -1,3-glucose units semi-alive at maturity lose their nucleus and other organelles; retain mitochondria, E.R., & plastids Oriented end-to-end forming a tube-like continuum Conducting dissolved sugars from a sugar-rich source to a sugar-poor sink region of the plant Source regions leaves sugars are synthesized; mature storage organs sugars may be released by the hydrolysis of starch Sinks actively dividing cells, developing storage organs, reproductive organs 1. Sieve cells have only sieve areas on both end and side walls; primitive; in all nonflowering vascular plants 2. Sieve tube members have both sieve areas and sieve plates; found only in flowering plants (angiosperms) *Sieve plates 1/more sieve areas at the end wall junction of 2 sieve tube members Albuminous cells parenchyma cells associated w/ sieve cells; albuminous cells and sieve cells derived from different parent cells; load/unload sugars into the cavity of the sieve cells Companion cells parenchyma cells associated w/ sieve tube members; companion cells and sieve tube members derived form same parent cell; load/unload sugars into the cavity of sieve tube members Stele organization of xylem and phloem in the stem 1. Protostele stellar type; central solid cylinder of xylem and phloem 2. Plectostele modification of the protostele; xylem and phloem interdigitate Cortex largely parenchymatous tissue bet. the epidermis and vascular tissue Endodermis Special cylinder of cells in some stems and all roots Casparian strip present in all endodermal cells; band/ring of lignin and suberin that infiltrates the cell wall; waterimpermeable material that binds to the plasma membrane of the endodermal cells Plasma membrane differentially control solute transfer Endodermis selectively controls w/c compounds are/are not absorbed Iigo Atienza 2Biology-3
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Chapter 3: Evolution & Diversity of Green and Land Plants Chapter 4: Evolution & Diversity of Vascular Plants

BOT 201
Plant Systematics 2 Ed. Michael G. Simpson
Root Specialized plant organs that function in anchorage and absorption of water and minerals Found in all vascular plants except for the Psilotales, Salviniales, etc. lost roots secondarily Develop by the formation of new cells w/in the growing apical meristem of the root tip Root tip region of continuous mitotic divisions Expansion both horizontally and vertically pushed the apical meristem tissue downward Single apical cell ancestral apical meristem of roots; in Selaginellaceae, Monilophytes Group of continuously dividing cells complex apical meristem; in Lycopodiaceae, Isoetaceae, seed plants 1. Rootcap/Calyptra covers the apical meristem; protects the apical meristem from mechanical damage; provides lubrication as the outer cells slough off 2. Root hairs hairlike extensions developed by epidermal cells away from the root tip; increase the surface area for absorption 3. Central vascular cylinder *pith center of CVS, if vasculature is lacking 4. Endodermis w/ casparian strips surrounds the vascular cylinder * Pericycle undifferentiated layer internal to the endodermis 5. Endogenous lateral roots in w/c new lateral roots originate by means of actively growing meristems, arising at the pericycle/endodermis *lateral roots penetrate the tissues of the cortex before exiting to the outside Mycorrhizae symbiotic relationship of roots w/ fungi o Fungal component aid the plant in increasing surface area for absorption and increasing the efficiency of selective mineral absorption; benefits in obtaining photosynthates from the plant Features used to classify vascular plants 1. Sporophyte vegetative morphology 2. Life cycle and reproductive morphology 3. Gametophyte morphology 4. Spore morphology Spores Laesura differentially thickened wall region corresponding to the tetrad attachment scar on each of the 4 immature spores ff. meiosis Spore types: a. Trilete spores w/ a 3-branched laesura b. Monolete spores w/ a linear, unbranched laesura c. Alete spores that lack laesura Rhyniophytes Paraphyletic first land plants w/ branched sporophytic axes Rhyniophyte sporophytes dichotomously branching axes bearing terminal sporangia that dehisce longitudinally Lacked roots and leaf-bearing shoot system Stems protostelic first-formed xylem centrarch Lycodiophyta Diverged after rhyniophytes Apomorphies: a. Dichopodial roots root apical meristem may branch into 2; no lateral roots developed Roots w/ endarch protoxylem protoxylem forms in a position interior to the metaxylem st *protoxylem 1 tracheary cells that develop w/in a patch of xylem; smaller, thinner that metaxylem c. Stems w/ exarch protoxylem protoxylem develops in a position exterior to the metaxylem d. Dorsiventral sporangia that dehisce transversely e. Sporophytic leaves dorsiventral organ; primary site for photosynthesis Lycophylls sporophytic leaves of lycophytes; single, unbranched vein, lacks a gap in the vasculature of the stem; develops by an intercalary meristem; develops from a shoot apex; originated from the transformation of enations Shoot stem and associated leaves Enations external, peg-like appendages that lack vascular tissue Lycopodiophyta Lycopodiopsida Lycopodiaceae Isoetopsida Selaginellaceae Isoetaceae 1. Lycopodiopsida (club mosses) Homospory 1 type of spore Long-lived sporophytic phase Sprorangia develop laterally in the axils of sporophylls Sporophylls specialized leaf that bears 1/more sporangia Strobilus/Cone terminal aggregate of sporophylls w/ associated sporangia a. Lycopodiaceae (club moss family) o Terrestrial or epiphytic (plant that grows on other plants) o Pendulous (apical), perennial herbs o Roots adventitious and endogenous; dichotomously branched (dichopodial); grow from the underground portions of the stem o Stems dichotomously branched rhizomes or corms; protostelic vasculature o Gemmae specialized short shoots; vegetative propagules o Leaves simple, sessile, spiral/whorled; blades scale-like/acicular, heteromorpic; w/ single midrib; ligule absent o Sporangia homosporous; reniform; occurring on short stalks in axils of sporophylls; globose/tetrahedral w/ trilete laesura o Sporophylls [photosynthetic, dispersed among vegetative leaves] or [ scale-like, nonphotosynthetic, organized in terminal strobili] o Gametophytes mycorrhizal; [epiterranean, photosynthetic] or [subterranean, saprophytic] o 5 genera: Huperiza, Lycopodiella, Lycopodium, Pseudolycopodiella, Phylloglossum o Worldwide distribution 2. Isoetopsida Possess leaf ligules b.
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Chapter 3: Evolution & Diversity of Green and Land Plants Chapter 4: Evolution & Diversity of Vascular Plants

Iigo Atienza 2Biology-3

BOT 201
Plant Systematics 2 Ed. Michael G. Simpson
*ligules tiny appendages on the upper (adaxial) side of the leaf, near the leaf base; providing hydration for developing lycophylls Heterosporous 2 types of spores; microspores and megaspores Microspores develop in microsporangia; small, produced in large numbers; forms into male gametophyte produces only antheridia spermmanufacturing organ Megaspores develop in megasporangia; larger, in fewer numbers; develops into a female gametophyte contains only archegonia housing the egg cell Both spore types may be produced in the same shoot or on different shoots Endosporic gametophytes gametes develop entirely w/in the original spore wall a. Selaginellaceae (Spike-Moss family) o Perennial herbs o Roots adventitious; dichotomous; from branch junctions, growing downward o Stems dichotomous; erect, cespitose, prostrate/repand, or climbing habit; pseudomonopodial or sympodial o Leaves simple, sessile, spiral; w/ single midrib; adaxially ligulate; blades small, homomorphic or dimorphic; 4 rows (2 dorsal rows smaller, 2 ventral/lateral rows larger) o Sporangia heterosporous; o Microsporangia, megasporangia occur on short stalks in the axils of ligulate sporophylls microsporophylls, megasporophylls o Gametophytes endosporic o Tropical and warm regions, worldwide b. Isoetaceae (Quillwort family) o Aquatic or terrestrial o Perennial herbs o Roots adventitious; endogenous from lower grooves in the stem; dichotomous o Stems protostelic; vertically oriented; cormlike; apical and lateral meristem lobed at the base o Leaves simple, spiral, in a basal rosette; blades widened, sheating, single midrib, ligulate at the apex of the expanded base o Sporangia heterosporous; adaxial side of sporophyll bases o Megasporangia outer leaves of a flush growth o Megaspores large, trilete, spore sculpturing; covered by velum; open by tissue degradation o Microsporangia inner leaves o Microspores small, monolete, numerous; covered by velum; open by tissue degradation o Gametophytes endosporic o Worldwide distribution Euphyllophyta Sister taxa of lycophytes Includes all other vascular plants Apomorphies: a. Monopodial roots they do not dichotomously branch at the apical meristem *lateral roots arise endogenously from the endodermis/pericycle Roots w/ exarch protoxylem protoxylem is positioned outer to the metaxylem c. Ancestral sporangia terminal in position w/ longitudinal dehiscence d. 30-kilobase inversion in the large single-copy region of chloroplast DNA e. Euphylls (=megaphyll) leaves of euphyllophytes; dorsiventral organs; primary site of photosynthesis; associated w/ leaf gap; highly branched system of veins; grow by apical/marginal meristems Leaf gap region of nonvascular, parenchyma tissue interrupting the vasculature of the stem Mesophyll chloroplast-containing tissue Trimerophytes common ancestor of lycophytes and euphyllophytes Planation flattening of the axes into a 2-dimmensional plane; axes - veins Webbing devt of thin tissue bet. the axes of the branches; web photosynthetic mesophyll Shoot stem and associated leaves; develop from the apical meristem Apical meristem o 1/more ultimate cells of the apical meristem undergo sequential mitotic cell divisions proliferation of cell derivatives o May contain 1 dominant apical cell, or a complex of several, actively dividing cells o 1/more of the derivatives maintain the position and function of the apical meristem o Cell derivatives undergo considerable elongation Leaf primordium matures into a leaf Node point of attachment a leaf to the stem Internode region bet. nodes Leaves o Upper and lower epidermis o Cuticle covers the outer cell wall of the epidermal cells o Veins contain xylem, phloem conduct water and sugars to/from the mesophyll cells Mesophyll a. Upper, columnar palisade mesophyll cells b. Lower, irregularly shaped spongy mesophyll cells; w/ large intercellular spaces Stomata gas exchange; lower epidermis Axil tissue at/above the region of junction of stem and upper leaf Bud immature shoot system; may develop into a lateral branch, or may terminate by developing into a reproductive structure Monopodial pattern of growth; lateral branches develop from axillary buds Euphyllophyta Monilophyta (Ferns) Equisetopsida (Horsetails) Equisetaceae Psilotopsida Ophioglossaceae (Ophioglossoid Ferns) Psilotaceae (Whisk Ferns) Marattiopsida (Marattoid Ferns) Marattiaceae Polypodiopsida/Leptosporangiatae Osmundales Osmundaceae b.
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Chapter 3: Evolution & Diversity of Green and Land Plants Chapter 4: Evolution & Diversity of Vascular Plants

Iigo Atienza 2Biology-3

BOT 201
Plant Systematics 2 Ed. Michael G. Simpson
Hymenophyllales Hymenophyllaceae Gleicheniales Gleicheniaceae Schizaeales Lygodiaceae Salviniales Marsileaceae Salviniaceae Cyatheales Cyatheaceae Polypodiales Aspleniaceae Dryopteridaceae Polypodiaceae Pteridaceae Monilophyta Monophyletic group Apomorphies: a. Siphonostele type of stem vasculature; a ring of xylem is surrounded by an outer layer of phloem (ectophloic) or by an outer and inner layer of phloem (amphiphloic/solenostele); central parenchymatous pith b. Stem protoxylem mesarch in postion tracheary elements first mature near the middle patch of xylem *protoxylem restricted to the lobes of the xylem 1. Equisetopsida (Horsetails) Equiseptophytes, sphenophytes, sphenopsids Monophyletic group Apomorphies: Ridged stems w/ internal hollow canal Reduced, whorled leaves marginally fused Sporangiophores consists of a peltate axis bearing longitudinally dehiscent sporangia Photosynthetic spores w/ elater Equisetum only extant species a. Equisetaceae Horsetail/Scouring-Rush family o Perennial herbs o Underground stems rhizomes o Aerial stems ridged; photosynthetic; w/ epidermis containing silica inclusions o Stem hollow pith, 2 rings of canals (lacunae) inner ring of carinal canals, outer ring of vallecular canals o Lateral branches develop from buds forming just above the nodal regions o Dimorphic aerial shoots few taxa; photosynthetic vegetative shoots and nonphotosynthetic reproductive shoots o Leaves small, simple, whorled, microphyllous (1-veined), nonphotosynthetic at maturity o Sporangia homosporous; in terminal strobili o Strobili an axis bearing numerous, peltate sporangiophores, each bearing 5-10 sporangia beneath the distal, hexagonal outer portion; longitudinal dehiscence o Spores lack an attachment scar (alete); spherical, green, each bearing 4 spatulate, hygroscopic elaters spore dispersal o Gametophytes photosynthetic; cushion-like o Worldwide distribution Subgenus Equisetum horsetails; w/ lateral branches o Subgenus Hippochaete scouring-rushes; w/o lateral branches Psilotopsida Apomorphies: Roots of ophioglossales lack root branches and root hairs Gametophytes nonphotosynthetic (heterotrophic); contain mycorrhizal fungi; subterranean a. Ophioglossales (Ophioglossoid Ferns) o Each leaf sterile segment and fertile segment o Sterile segment contains the photosynthetic blade/lamina o Fertile segment bears the sporangia o Underground rhizome unbranched roots that lack root hairs o Eusporangia sporangia of ophioglossales and land plants except leptosporangia; large, derived from epidermal cells; sporangial wall more than 1 cell layer; ancestral condition Ophioglossaceae (Adders Tongue family) - Terrestrial, perennial herbs - Roots fleshy, mycorrhizal, lack root hairs - Stems subterranean, erect; vasculature protostele/ectophloic siphonostele - Leaf solitary, lacking circinate vernation; blade simple, unlobed/compound - Sori lacking - Sporangia homosporous, eusporangiate; on stalked fertile segment; transverse dehiscence - Spores tetrahedral, trilete - Gametophytes nonphotosynthetic; mycorrhizal/mycotrophic - Worldwide distribution b. Psilotales (Whisk Ferns) o 2 genera Psilotum, Tmesipteris o Sporophyte independent, dominant, free-living; horizontal rhizome aerial stems o Gametophyte small, obscure, free-living o Epiphytic; w/ mycorrhizal rhizomes o Roots lacking; lost secondarily o Leaves reduced, peg-like o Enations leaf that lacks vascular tissue o Sporangia synangium 2/3 lobed; fusion product of 2/3 sporangia o Gametophytes nonphotosynthetic o Tropical regions Psilotaceae - Terrestrial, epiphytic - Perennial herbs - Roots absent - Underground stems rhizoid-bearing rhizomes w/ endophytic mycorrhizae - Aerial stems photosynthetic; terete, ridged, or flattened - Dichotomous - Vasculature protostele/solenostele - Leaves simple, spiral, distichous; blades small - Sporangia eusporangiate, homosporous, arising from short, lateral branches o
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Chapter 3: Evolution & Diversity of Green and Land Plants Chapter 4: Evolution & Diversity of Vascular Plants

Iigo Atienza 2Biology-3

BOT 201
Plant Systematics 2 Ed. Michael G. Simpson
- Synangiate 3(Psilotum) or 2(Tmesipteris) sporangia - Spores reniform, monolete - Gametophytes nonphotosynthetic, cylindrical, saprophytic, mycorrhizal - Sperm multiflagellate - Tropical or warm temperate regions Marattiopsida (Marattoid Ferns) 6 genera Large pinnate/bipinnate leaves Circinate vernation Sporangia abaxial (ventral) surface of leaflet blades Gametophyte photosynthetic Synangium sporangia are fused into a common structure a. Marattiaceae o Large, terrestrial ferns o Tissues w/ mucilage canals o Stems rhizomatous/erect; stout and trunk-like; polycyclic dictyostele o Leaves circinate vernation; mature large; stipulate, simple or 1-4 pinnate; petioles and rachillae w/ swollen pulvini and prominent pneumatodes o Sori abaxial, intramarginal, exindusiate o Sporangia eusporangiate, homosporous, vertically dehiscent o Spores trilete/monolete/alete o Gametophyte large, thalloid, photosynthetic Polypodiopsida/Leptosporangiatae Sporophytes perennial herbs or trees; may be annuals Rhizome horizontally-oriented stem; most leptosporangiate ferns; grow under/upon the ground Terrestrial, aquatic, epiphytic Arborescent some L. ferns; tall, erect, aerial stem Vines few L. ferns; w/ weak stems or w/ elongate vine-like leaves that sprawl on the ground or upon another plant Stem: [ectophloic or amphiphloic siphonostele/solenostele], dictyostele, or protostele Circinate vernation immature leaves; both major and minor axes or leaf divisions are coiled early in devt and uncoil at maturity Fiddleheads/crosiers young, coiled leaves Frond leaf Stipe petiole st Pinnae 1 discrete leaflets or blade divisions of a fern leaf Pinnules/segments ultimate leaflets or blade divisions Leaves dimorphic fertile leaves, sterile leaves Leaf venation pinnate central vein giving rise to veinlets on either side; palmate less common, more than 1 main vein arising from the base a. Open (free) veins arising from the midvein/base of a pinnule do not join back together; simple, forked, or bifurcate b. Reticulate (anastomosing) veins appear to join back together, enclosing an area (areole) Trichomes (hair-like structures) or scales Clathrate scales w/ the cell walls of adjacent cells that are thick Non-clathrate w/ thin anticlinal cell walls Leptosporangium develops from a single cell; single layer of cells making up the sporangial wall; smaller spore number; proximal stalk, distal sporangial body *devt: gradate (mature in succession), simultaneous (mature at the same time), mixed, or intermingled (no pattern) Annulus part of the wall of the sporangial body develops into a single row of specialized cells Sori aggregation of discrete leptosporangial clusters Receptacle point of attachment of leptosporangia w/in the sorus Paraphyses sterile, hair-like structures Indusium flap of tissue arising from the blade surface that covers the sori; protect leptosporangia, regulates spore dispersal *reniform, orbicular, or linear *peltate, or lateral attachment of indusium Acrostichoid leptosporangia not aggregated into definable sori False indusium reflexed extension of the blade margin; overlaps the sorus Perine outer wall layer of the spore Gametophyte superficial, w/ rhizoids; bear antheridia, archegonia *sperm cells coiled, multiflagellate Osmundales Osmundaceae (Cinnamon Fern family) o Terrestrial o Stem erect; ectophloic siphonostele o Leaves 1-2 pinnate or pinnatifid; stipulate at the base of petiole; dimorphic [fertile and sterile leaves] or w/ [fertile and sterile leaf segments] o Sori and indusial absent o Sporangia large bodies, short stalks; dehisce by apical slit (annulus lateral) o Spores green, subglobose, trilete; 128-512 per sporangium o Gametophytes large, green, cordate, superficial o Chromosome no. = 22 o Tropical, temperate regions Hymenophyllales Hymenophyllaceae (Filmy Fern family) o Epiphytic, terrestrial herbs o Stems rhizomatous; rhizomes w/o scales; protostelic; [slender and creeping] or [stout and erect] o Leaves 1-cell thick; stomata absent; cuticle absent/reduced; blade scales absent; trichomes sometimes present; venation open o Sori marginal o Receptacle elongate, continuous w/ vein tips o Indusia present; conicular, tubular, or 2-lobed (bivalvate) o Sporangia basipetalous o Annulus oblique o Spores green, globose, trilete o Gametophytes filamentous/ribbon-like; reproduce by fragmentation/gemmae
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Chapter 3: Evolution & Diversity of Green and Land Plants Chapter 4: Evolution & Diversity of Vascular Plants

Iigo Atienza 2Biology-3

BOT 201
Plant Systematics 2 Ed. Michael G. Simpson
o Pantropical, south-temperate regions Gleicheniales Gleicheniaceae (Forking Fern family) o Terrestrial; in dense, open thickets o Stems rhizomatous creeping, dichotomous, w/ a vitalized protostele o Leaves indeterminate growth; rachises pseudodichotomous; 1-2 pinnate, free veins o Sori round, abaxial, not marginal, exindusiate o Sporangia round to pear-shaped; 5-15 sporangia per sorus develop simulatenously o Annulus transverse o Spores globose-tetrahedral or bilateral; trilete/monolete; 128-800 per sporangium o Gametophytes large, green, superficial, w/ club-shaped hairs; endotrophic mycorrhizae o Pantropical Schizaeales Lygodiaceae o Terrestrial, climbing o Stems rhizomatous; rhizomes slender, creeping, bearing hairs; protostelic o Leaves indeterminate w/ an elongate, twinning and climbing rachis that bears pinnae alternately o Pinnae pseudodichotomous; veins free/anastomosing o Sori abaxial; on lobes of ultimate leaf segments o Sporangia 1 per sorus o Annulus transverse, subapical, continuous o Spores tetrahedral, trilete; 128-256 per sporangium o Gametophytes green, cordate, superficial o Chromosome no. = 29, 30 o Pantropical Salviniales Aquatic Sporangia heterosporous 2 types of spores and sporangia (megaspores in megasporangia; microspores in microsporangia); form w/in a sporocarp Sporocarp rounded, seed-like, structure w/ a hard outer layer; protection, dormancy Endospory devt of gametophyte w/in the original spore wall Marsileaceae (Clover Fern family) o Rooted, aquatic herbs o Stems elongate, slender, creeping rhizomes; w/ aerenchyma; solenostele o Leaves circinate, simple, or palmate w/ 2/4 sessile leaflets; veins dichotomous o Sporocarps reniform w/ a stalk arising from the petiole base or leaf axil o Sori consists of a column of megasporangia and microsporangia enveloped by hood-like indusium; lack annulus o Sorophore elongate, gelatinous structure; w/ several pairs of sori attached o Megasporangium bears 1, trilete megaspore after imbibing in water produce acrolamellae o Acrolamellae w/ apical, longitudinal folds and basal horizontal folds; contain sperm lake o o Sperm lake central, liquid filled region Microsporangia produce several, trilete microspores o Subcosmopolitan Salviniaceae (Floating Fern family) o Floating, aquatic herbs o Roots absent in Salvinia, present in Azolla o Stems dichotomously branching rhizomes; protostelic; aerenchymatous o Leaves simple, dimorphic, aerenchymatous, distichous at maturity; blades round-oblong o Sporocarp globose, heterosporous; bear [1 megasporangium] or [several microsporangia] o Megasporangium 1 functional megaspore surrounded by massulae form female gametophyte w/ several protruding archegonia *massulae gelatinous masses of tissue from multinucleate plasmodium o Microsporangia bear several microspores o Chromosome no. = 9 (Salvinia), 22 (Azolla) o subcosmopolitan Cyatheales Cyatheaceae (Scaly Tree Fern family) o Mostly terrestrial, some epiphytic o Stems arborescent; polycyclic dictyostele o Trunks w/ marescent leaves or leaf bases o Shoot apices and petiole bases covered w/ large scales/hairs o Leaves large; blades 1-3 pinnate; petioles w/ discontinuous pneumathodes in 2 lines; blade veins free, simple-forked o Sori abaxial, round, superficial/terminal on veins; marginal/submarginal; receptacle raised; paraphyses present; exindusiate/indusiate o Indusium saucer-like, cup-like, bivalvate, globose; surround completely sporangia o Sporangia mature gradately o Annulus oblique o Spores tetrahedral, trilete, variously ornamented o Gametophytes green, cordate o Chromosome no. = 69 o Pantropical Polypodiales Aspleniaceae (Spleenwort family) o Terrestrial, epipetric, or epiphytic perennials o Stems rhizomatous o Rhizomes creeping, climbing, ascending, or suberect; bear clathrate scales at shoot apices and petiole bases o Leaves monomorphic; simple to multipinnate; w/ small clavate hairs; 2 back-to-back C-shaped vascular strands at petiole base o Sori, indusial elongate along veins o Sporangia mixed; sporangial stalks of 1 row, long o Spores reniform, monolete, w/ winged perine o Chromosome no. = 36, rarely 38,39 o subcosmopolitan Dryopteridaceae o Terrestrial, epipetric, epiphytic perennials o Stems rhizomatous
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Chapter 3: Evolution & Diversity of Green and Land Plants Chapter 4: Evolution & Diversity of Vascular Plants

Iigo Atienza 2Biology-3

BOT 201
Plant Systematics 2 Ed. Michael G. Simpson
o Rhizomes dictostelic; creeping, ascending, erect, or scandescent-climbing o Shoot apices w/ non-clathrate scales o Leaves monomorphic, rarely dimorphic; simple/pinnate to decompounds; veins pinnate/forking, free to anastomosing, w/ or w/o included veinlets o Sori round; indusiate/exindusiate; acrostichoid in some o Indusia round-reniform or peltate o Sporangia mixed; sporangial stalks in 3 rows o Spores reniform, monolete o Pantropical, temperate o Chromosome no. = 41 Polypodiaceae o Epiphytic, epipetric, or terrestrial perennials o Stems rhizomatous o Rhizomes dictyostelic, long-short creeping, bear scales o Leaves simple to 1-pinnate; monomorphic/dimorphic; blades glabrous or w/ hairs/scales; veins anastomosing/reticulate o Sori abaxial, round, oblong, or elliptic; rarely elongate/acrostichoid; receptacle w/ paraphyses; exindusiate o Sporangia mixed, sporangial stalks in 1-3 rows o Spores hyaline to yellowish; reniform; monolete o Chromosome no. = 35, 36, 37 o Pantropical, temperate Pteridaceae o Terrestrial, epipetric, epiphytic; rarely aquatic o Stem rhizomatous o Rhizomes creeping to erect; bear scales/hairs o Leaves simple, pinnate, pedate, or decompound; veins free/anastomosing o Sori exindusiate; [marginal w/ false indusium] or [intramarginal in lines along veins]; receptacle not raised o Sporangia mixed; sporangial stalks 1-3 cells thick o Spores globose or tetrahedral, trilete, ornamented o Chromosome no. = 29, 30 o Subcosmopolitan; tropical and arid region
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Chapter 3: Evolution & Diversity of Green and Land Plants Chapter 4: Evolution & Diversity of Vascular Plants

Iigo Atienza 2Biology-3

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