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Vascular Plants
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Generalized life cycle
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Moss life cycle
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Spore -> Gametophyte
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Gametophyte -> Zygote
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Zygote -> mature sporophyte
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Sporophyte -> spores
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Vascular Plants
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Vascular Plants Branching sporophyte in Silurian
First vascular plants in Devonian First flowering plants Age of Ferns First fossil of non-algal land plant in Silurian mya Liverwort spore tetrads - end of Ordovician
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Fossils Probable embryophyte spores at 450 Ma
Cooksonia fossils in Silurian (ca. 430 Ma)
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Plants like Cooksonia lacked a vascular system
Dichotomously-branching axes Terminal sporangia No roots or leaves
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Over-time became larger, more complex, and acquired a vascular system
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Living vascular plants (Tracheophytes)
Stems and roots (often leaves)
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Living vascular plants (Tracheophytes)
Stems and roots Sporophytes dominate the life-cycle
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Sporophyte dominance Gametphyte Tree fern
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Why sporophyte dominance?
Spore dispersal by wind: aided by height Competition for light (gametophyte constrained by the need for water)
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Living vascular plants (Tracheophytes)
Stems and roots Sporophytes dominate the life-cycle A vascular (transport) system
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Vascular system Xylem (water transport) and phloem (metabolite transport) Stem Vascular Bundle Xylem Phloem Root
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Xylem Composed primarily of Tracheids Elongated, dead, cells
Cell wall impregnated with lignin Transport of water from soil to leaves
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Phloem Transport via sieve elements Elongated, living cells
Transport of sugars, hormones, etc.
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3 Major groups of Vascular plants
Seed plants (gymnosperms and angiosperms) Lycophytes (club mosses and their relatives) Moniliforms (ferns and fern allies)
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Major tracheophyte taxa
Seed Plants (ca. 290,000 species) Lycophytes (ca. 1,100 species) Ferns and allies (ca. 11,000 species) Tracheophytes that are not seed plants are sometimes called “pteridophytes”
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Vascular plant phylogeny
bryophytes lycophytes Ferns + Seed plants Microphylls Cooksonia Megaphylls Vascular tissue, Roots Branched sporophyte
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Organ systems origins Stems - dichotomous branching
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Organ systems origins Stems Leaves Microphylls
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Organ systems origins Stems Leaves Microphylls Megaphylls
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Organ systems origins Stems Leaves Microphylls Megaphylls Roots
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Homospory versus Heterospory
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An important variation: Heterospory
Megasporangium Microsporangium Lycopodium (homosporous) Selaginella (heterosporous)
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Heterospory Microspores and megaspores produced in different sporangia on different leaves (microsporophylls; megasporophylls) Microspores grow into male gametophytes Megaspores grow into female gametophytes - remains within spore wall
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Homospory Haploid Diploid sperm zygote egg sporophyte gametophyte
spore
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Heterospory Haploid Diploid zygote egg sperm sporophyte microspore
female gametophyte male gametophyte microspore megaspore
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Heterospory evolved many times. Why?
Increases potential for outcrossing Specialization of function between micro- and megagametophyte permits greater efficiency (less cost)
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3 Major groups of Vascular plants
Seed plants (gymnosperms and angiosperms) Lycophytes (club mosses and their relatives) Moniliforms (ferns and fern allies)
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Moniliforms Spermatophyta Lycophyta Pryer et al. 2001
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Lycophytes 380 Ma old 1100 spp. Microphylls only
Sister group to the other living vascular plants
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Lycophytes 380 Ma old 1100 spp. Microphylls only lycophytes Ferns +
Seed plants Microphylls Megaphylls
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Lycophytes Lycopodium Selaginella Clubmoss (Lycopodium) Isoetes
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Carboniferous lycopods up to 40 m
Sigillaria Lepidodendron
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Lycopodium life cycle I
Sporophyte makes sporangia often in a “strobilus” Sporangium
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Lycopodium life cycle I
Sporophyte makes sporangia often in a “strobilus” Sporophyll Sporangium
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Lycopodium life cycle II
Spores dispersed by wind: germinate into a minute gametophyte Spore Thallus Spore Rhizoids
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Lycopodium life cycle III
Gametophyte produces archegonia and antheridia (bisexual) Biflagellate sperm fertilize egg cells New sporophyte grows Archegonial neck
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Lycophyte diversity 3 Major groups Lycopodiaceae (club mosses)
Selaginella Isoetes
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Lycopiaceae Approximately 400 species
Dominated Carboniferous, up to 40 m tall form much of modern coal Homosporous Archegonia and Antheridia can take 6-15 years to mature
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Selaginella Approximately 700 extant species Heterosporous
Moist habitats or "resurrect" Selaginella umbrosa
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Isoetes Approximately 200 species Grow in water or dried pools
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Moniliforms Spermatophyta Lycophyta
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Ferns and fern allies (moniliforms)
Includes ferns (Pterophyta) and two small groups (Psilophyta and Sphenophyta) ca. 12,000 spp. Homosporous or heterosporous Megaphylls (lost in Psilophyta and Sphenophyta) Ecologically important especially as tropical epiphytes
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Moniliforms Ferns and Fern Allies
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Ferns and Fern Allies Equisetum Psilotum
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Ferns and Fern Allies eusporangiate ferns Sporangium wall has 2 or more cell layers
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Ferns and Fern Allies leptosporangiate ferns Sporangium wall has 1 cell layer
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Ferns and Fern Allies Psilotum and Ophioglossum
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Psilotaceae/Psilophyta
Psilotum (2 spp.) and Tmesipteris (15 spp.) No roots and reduced or absent leaves, photosynthetic stems Sporangia on lateral branches Homosporous Thought to be "primitive vascular plants", but more likely simplified due to association with fungi.
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Psilotaceae Psilotum Sporangium Tmesipteris Psilotum
Reduced forked leaves Psilotum
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Psilotum Dichotomizing stem, no roots Long-lived gametophytes
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Ophioglossaceae Adder's tongue, eusporangiate ferns
Homosporous Worldwide, common in disturbed areas Botrychium (~60 spp.) and Ophioglossum (30 spp.) Ophioglossum can have upwards of 1400 chromosomes - perhaps more then any other organism
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spore bearing sporophore
Ophioglossaceae Sterile blade
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Botrychium virginianum
Ophioglossaceae
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Ferns and Fern Allies Equisetum
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Equisetaceae Horse tails
Equisetum (15 spp.) Homosporous Dates back to Devonian, with 20 m high stems - lots of diversity in Carboniferous forests Extant species "living fossils" Leaves whorled, fused into sheaths at base, only microphylls
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Equisetum
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Extinct trees Calamites (Carboniferous) Calamites
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Equisetum gametophyte (hermaphroditic)
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leptosporangiate ferns
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Osmunda Most ferns species are Leptosporangiate Ferns
Large megaphylls (fronds) unfold lengthwise from a "fiddlehead" Osmunda
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Section through sorus Sporangium Sporangia are arranged in sori Sori
Indusium Sporangia Sori Section through sorus Sporangia are arranged in sori Sporangium
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Fern gametophyte (prothallus)
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Ferns Tree ferns (Cyatheaceae) Maidenhair fern Walking fern
Epiphytic fern (Platycerium)
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Leptosporangiate ferns
App. 11,000 species in recognized groups (most of fern diversity) Cover 4 of the major clades Marsileaceae Osmundaceae Cyatheaceae Polypodiaceae
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Marsileaceae Salvinia Marsilea Mostly aquatic
Leave blade divided into 2-4 leaflets (clover-like) Heterosporous - megagametophytes with only one archegonium Symbiotic with cyanobacteria, fertilized rice fields Spores remain viable for a century Salvinia Marsilea
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Osmundaceae Sporangia loose, not in sori Homosporous Osmunda Todea
Leptopteris
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Cyatheaceae Tree fern growth Sporangia in sori on bottom of leaf
Stem usually single and erect Alsophila Cyathea
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Polypodiaceae Asplenium Nephrolepis Elaphoglossum Adiantum Polypodium
Pteris Tectaria
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Main points Features of vascular plants Homospory versus heterospory
Megaphylls vs. microphylls Life cycle of the fern Fern allies: Psilotum, Equisetum
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