Chapter 4: Evolution and Diversity of Vascular Plants

Michael G. Simpson

Vascular Plants = Tracheophyta (Tracheophytes)

Apomorphies?

Alternation of Generations: Haploid (n) & Diploid (2n) adult phases

Bryophytes: Gametophyte dominant, long-lived

Bryophytes (Liverworts, Hornworts, Mosses): Gametophytes are dominant, long-lived

Alternation of Generations: Haploid (n) & Diploid (2n) adult phases

Vascular Plants: Sporophyte dominant, long-lived

Vascular Plants: Sporophytes are dominant

Sporophytic axes branched with multiple sporangia

Rhynia

Two early branching patterns in vascular plants:

lignin - hard substance secreted within secondary cell wall Major adaptive: added structural support enabled vascular plants to grow much larger.

secondary cell wall

- forms between 1 cell wall and plasma membrane in some plant cells - adds structural support - found in tracheids, vessels, fibers (all dead cells)

lignified secondary cell wall

Sclerenchyma 1) Secondary cell wall (+ primary) 2) Dead at maturity (usually)

Fibers Elongate, sharply tapering

Sclerenchyma 1) Secondary cell wall (+ primary) 2) Dead at maturity (usually) Sclereids Isodiametric to irregular

Tracheary elements 1) cells that function in water/mineral conduction 2) cells dead at maturity, with lignified, 2 cell walls 3) cells arranged end-to-end, forming long tubes Xylem = tissue composed of: 1) tracheary elements 2) parenchyma 3) fibers (These have common origin/location/function.)

Tracheary Elements
Vessels perforation plates = holes in end walls. Tracheids imperforate (only pits at end walls). Found in most monilophytes (except Evolved independently from preexisting Equisetum and some lepto. ferns) and most tracheids in Equisetum, a few leptosporangiate ferns, all Gnetales, and almost all angiosperms gymnosperms (except Gnetales)

Sieve Elements
-cells with sieve areas/plates, having pores lined by callose callose (polysaccharide composed of -1,3-glucose units) -lines pores of sieve areas! Sieve cells only sieve areas All vasc. pls. except angiosperms Sieve tube members - Sieve plates Apomorphy of Angiosperms

Phloem
= tissue composed of: 1) sieve elements 2) parenchyma 3) fibers These have common origin/function.

Endodermis single layer of cells surrounding vasculature of roots and some underground stems.

Casparian Strip: forces fluids from outside through plasma membrane = selective absorption

Rhynia - one of earliest vascular plants

(ca. 400 million years ago)

- lacked roots

Roots
Function in anchoring and absorption.

Root

apical meristem - region of actively dividing cells

Roots
Five diagnostic features: 1) Root cap functions to protect apical meristem and lubricate root during growth. 2) Root hairs function to greatly increase water/mineral absorption (exception Psilotopsida). 3) Central vascular cylinder site of conduction of water/minerals and sugars. 4) Endodermis Casparian strip functions in selective absorption. 5) Endogenous lateral roots develop from pericycle or endodermis, penetrate to outside.

Mycorrhizae
Symbiotic association with various species of fungi. Found in many, if not most, vascular plants.
Fungus aids plant by 1) increasing surface area for water and mineral absorption 2) increasing efficiency of mineral absorption. Fungus benefits in obtaining photosynthates (sugars and other nutrients) from the plant.

Tracheophyta Classification

Lycopodiophyta lycophytes Apomorphies: 1)Roots dichopodial -apical meristem may branch into two. 2) Root protoxylem endarch (to inside) Protoxylem first xylem formed Metaxylem later xylem

3) Stem protoxylem exarch (to outside)

4) Sporangia dorsiventral (flattened upper and lower surfaces) with transverse dehiscence.

Shoot = stem + leaves

Leaf sporophytic

Lycophyll simple leaf with 1 vein, no leaf gaps.

Tracheophyta Classification

LYCOPODIOPSIDA Lycopodiaceae
homosporous = 1 type of spore 5 genera: Huperzia, Lycopodiella, Lycopodium, Pseudolycopodiella, Phylloglossum)

Lycopodium digitatum

ISOETOPSIDA Selaginellaceae + Isoetaceae

1) Leaves ligulate 2) Heterosporous 2 types of spores

Ligule tiny appendages on upper side of leaf (near base)

Heterospory two types of spores: megaspores (from megasporangia) microspores (from microsporangia)

Isoetaceae Quillwort family

1 genus: Isoetes (ca. 200 spp.)

Isoetes quillwort/Merlins grass

Lycopods native to San Diego area:

Isoetes Merlins Grass

Isoetes orcuttii Isoetes howellii

Selaginellaceae Spike-moss family

1 genus: Sellaginella (700 spp.)

Selaginella spike-moss

Selaginella bigelovii

Selaginella cinerascens

Some Selaginella have dimorphic leaves

2 rows large leaves

Selaginella apoda

Some Selaginella have dimorphic leaves

2 rows small leaves

Selaginella apoda

Extinct Lycopods were tall trees: make up some of coal deposits

Lepidodendron

Euphyllophyta Apomorphies: 1) roots monopodial 2) root protoxylem exarch 3) sporangia terminal on lateral branches, longitudinally dehiscent

Archeopteris

Euphyllophyta Apomorphies: 1) roots monopodial 2) root protoxylem exarch 3) sporangia terminal on lateral branches, longitudinally dehiscent 4) shoot with euphylls shoot = stem + leaves

Euphyllophyta Apomorphies: 1) roots monopodial 2) root protoxylem exarch 3) sporangia terminal on lateral branches, longitudinally dehiscent 4) shoot with euphylls

Leaves of euphyllophytes: a)evolved by planation & webbing of stems b)have multiple veins c)have a leaf gap parenchyma replaces vascular tissue upper junction of leaf trace with stem vasculature

Euphyllophyta Apomorphies: 1) roots monopodial 2) root protoxylem exarch 3) sporangia terminal on lateral branches, longitudinally dehiscent 4) shoot with euphylls 5) 30 kb chloroplast DNA inversion

Tracheophyta Classification

Monilophyta -monilophytes
Apomorphies: 1) stem protoxylem mesarch 2) siphonostele

Equisetopsida (Equisetophytes) Equisetaceae Equisetum Horsetails

/ Scouring Rushes

Equisetopsida Apomorphies: 1) stem ribbed with canals 2) leaves reduced, whorled

Equisetopsida Apomorphies: 1) stem ribbed with canals 2) leaves reduced, whorled 3) sporangiophore 4) spores with elaters, chlorophyllous

cone (strobilus)

whorled microphylls

Equisetum hyemale common scouring-rush

Equisetum laevigatum smooth scouring-rush

Subgenus Hippochaete souring-rushes

Equisetum arvense Common Horsetail

Subgenus Equisetum horsetails

Calamites -fossil (extinct) equisetophyte, was tree-sized, makes up coal deposits today

Psilotopsida
Apomorphies: 1) Roots unbranched, root hairs absent. 2) Gametophyte subterranean, mycorrhizal.

Psilotopsida
Two members: 1) Ophioglossales ophioglossoid ferns 2) Psilotales whisk ferns

Ophioglossales: One family: Ophioglossaceae 4 genera, incl. Botrychium, Ophioglossum

Ophioglossaceae Ophioglossum californicum

Calif. Adders Tongue

Psilotales
Apomorphies: 1) Roots lost. 2) Leaves reduced (microphylls or enations). 3) Synangium with bifid appendage.

Psilotales
One family: Psilotaceae 2 genera: Psilotum Tmesipteris

dichotomous branching

Psilotum nudum

enations / microphylls

Psilotum nudum

synangium

bifid (2-forked) appendage

Psilotum nudum

Marattiopsida Marattioid Ferns

Apomorphy: 1) Polycyclic siphonostele

Marattiaceae

eusporangia

Polypodiopsida
Leptosporangiate Ferns Apomorphy: Leptosporangium

Polypodiopsida
Leptosporangium: - one cell thick - spores ejected

Stem: rhizome trunk vine

fern leaves can have specialized terminology: leaf = frond petiole = stipe major divisions = pinna (1, 2, etc.) ultimate divisions = pinnules

fern leaf venation: open (simple or forked), reticulate

fern scales: e.g., clathrate or non-clathrate

sorus (pl. sori) aggregation of sporangia indusium - flap

shape / morphology

leptosporangium / annulus -varies with different groups/families

sperm

egg

Fern leaf development: circinate

fiddle head - crozier

Salviniales
Apomorphies:
1) aquatic adaptation 2) heterospory (independently evolved) 3) sporocarps (dormant, seed-like body, house specialized sporangia)

Two famlies:
Marsileaceae clover fern family Salviniaceae floating fern family

Marsilea

Salvinia

Azolla

Marsileaceae

Marsilea Pilularia

Salviniaceae

Azolla

Salvinia

Azolla Mosquito Fern - symbiotic relationship with blue green bacteria; seeded in rice paddies for nitrogen fixation - inhibits mosquito population by covering surface - fodder for animals

Cyatheales: Cyatheaceae
largest family of tree ferns (trunk arborescent)

Polypodiales: A few families: Aspleniaceae

-linear sori / indusia

Cyathium a tree fern

Polypodiaceae
- sori exindusiate (naked)

Polypodiaceae: sorus
Polypodium californicum California Polypody

exindusiate

Polypodiaceae
Platycerium Staghorn Fern

sporangia acrostichoid (not aggregated into sori)

Pteridaceae
-false indusium

Cyathium a tree fern

Pteridaceae

Adiantum reniformis California Maidenhair Fern