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Home , Lycopodiopsida

Introduction to . Lecture 29

Alexey Shipunov

Minot State University

November 14, 2014

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 1 / 32 Outline

1 Questions and answers

2 diversity Pteridophyta

3 Plant diversity “Ferny”

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 2 / 32 Outline

1 Questions and answers

2 Plant diversity Pteridophyta

3 Plant diversity Heterospory “Ferny” ferns

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 2 / 32 Outline

1 Questions and answers

2 Plant diversity Pteridophyta

3 Plant diversity Heterospory “Ferny” ferns

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 2 / 32 Growth is not restricted Vascular tissues Endoderm and root system (not in all ferns)

Questions and answers Previous final question: the answer

Why are ferns more advanced than ?

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 3 / 32 Questions and answers Previous final question: the answer

Why are ferns more advanced than mosses?

Growth is not restricted Vascular tissues Endoderm and root system (not in all ferns)

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 3 / 32 Plant diversity Pteridophyta

Plant diversity Pteridophyta

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 4 / 32 Plant diversity Pteridophyta Pteridophyta: ferns and allies

≈ 12, 000 species and six classes Sporic life cycle with predominance is often reduced to prothallium (small -like plant), some Pteridophyta have male and female Have true roots (only whisk ferns, Psilotopsida are exception) Homoiohydric (same as seed plants) Sporophyte always starts development from embryo located on gametophyte Have true xylem and phloem, but do not have secondary thickening (exceptions: and extant Isoëtes and Botrychium)

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 5 / 32 Plant diversity Pteridophyta Pteridophyta classes

Subphylum Lycopodiophytina () Class Lycopodiopsida Subphylum Pteridophytina (monilophytes) Class Equisetopsida (horsetails) Class Psilotopsida (whisk ferns) Class Ophioglossopsida (ophioglossalean ferns) Class Marattiopsida (giant, or marattialean ferns) Class Pteridopsida (“true” ferns)

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 6 / 32 Plant diversity Pteridophyta Lycopodiopsida

Four main genera (, , and Isoëtes) and ≈ 1000 species Separate, microphyllous* lineage of Pteridophyta (all other groups are megaphyllous) Sporangia associated with and often form *. Spermatozoon typically with two flagella (like in mosses). Homosporous genera have achlorophyllous, mycoparasitic underground gametophyte. In the past, were dominant trees of tropical swamp forests (lepidodendrids) and their remains became a Two genera, Selaginella (spike ) and Isoëtes (quillwort) are heterosporous.

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 7 / 32 Plant diversity Pteridophyta Tropical , Huperzia linifolia

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 8 / 32 Plant diversity Pteridophyta drummondii, one of smallest lycophytes

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 9 / 32 Plant diversity Pteridophyta Before: Chicago 300 Million Years Ago (lepidodendrids)

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 10 / 32 Plant diversity Pteridophyta After: aquatic lycophyte Isoëtes sp.

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 11 / 32 Plant diversity Pteridophyta Equisetopsida

Small group of one genus, with ≈ 30 species Leaves are reduced into scales, stems are segmented, photosynthetic. Have specific stele—artrostele with specific central, valecular and carinal canals (similar to stele of some grasses) Sporangia associated with specialized leaves—sporangiophores. have attached elaters. Gametophyte minute, usually dioecious but plants are homosporous

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 12 / 32 Plant diversity Pteridophyta Strobili and sporangiophores of Equisetum arvense

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 13 / 32 Plant diversity Pteridophyta Equisetum giganteum

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 14 / 32 Plant diversity Pteridophyta Equisetum sp. elaters

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 15 / 32 Plant diversity Pteridophyta Artrostele

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 16 / 32 Plant diversity Pteridophyta Horsetail gametophytes

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 17 / 32 Plant diversity Heterospory

Plant diversity Heterospory

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 18 / 32 Plant diversity Heterospory Horsetails start it: spores same, gametophytes different

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 19 / 32 Plant diversity Heterospory Heterospory

Heterosporous ferns (lycophytes Selaginella and Isoëtes, monilophytes Salvinia, Marsilea, Pilularia, Regnellidium and Azolla) went one step further and made their spores different too. It will allow the better allocation of resources and will restrict the self-fertilization.

Terms covered: Male gametophyte, female gametophyte and microsporangium Megaspores and megasporangium

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 20 / 32 Plant diversity Heterospory Heterosporic cycle: differences

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 21 / 32 Plant diversity “Ferny” ferns

Plant diversity “Ferny” ferns

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 22 / 32 Plant diversity “Ferny” ferns Psilotopsida

Small tropical group of two genera, Psilotum and Tmesipteris and 7 species Have protostele (like lycophytes), underground long-lived gametophytes but multiflagellate spermatozoa (like horsetails and all ferns). Sporangia unite into synangia. Leaves may absent (Psilotum) and replaced with enatia. Externally remain , the oldest extinct Pteridophyta

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 23 / 32 Plant diversity “Ferny” ferns Hawaiian Psilotum complanatum

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 24 / 32 Plant diversity “Ferny” ferns New Zealand Tmesipteris tannensis with double synangium

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 25 / 32 Plant diversity “Ferny” ferns Ophioglossopsida

Small group (Ophioglossum, Botrychium, Mankyua and Helminthostachys) and ≈ 75 species Always have underground rhizome and aboveground bisected leaves: one half is the blade and other half is sporangiophore. Gametophytes grow underground Some (Botrychium, grape ) have secondary thickening of underground rhizome. Ophioglossum vulgatum, adder’s tongue fern, has 2n = 1360, the largest chromosome number ever.

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 26 / 32 Plant diversity “Ferny” ferns Ophiloglossum vulgatum, 2n = 1360 hero

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 27 / 32 Plant diversity “Ferny” ferns Helminthostachys zeylanicum (Ophioglossopsida)

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 28 / 32 Plant diversity “Ferny” ferns Mankyua chejuense (Ophioglossopsida)

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 29 / 32 Why does heterospory help ferns to be more adapted for the life on land?

Plant diversity “Ferny” ferns Final question (3 points)

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 30 / 32 Plant diversity “Ferny” ferns Final question (3 points)

Why does heterospory help ferns to be more adapted for the life on land?

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 30 / 32 Plant diversity “Ferny” ferns Summary

Heterosporous plants have two kinds of spores: female (megaspores) and male (microspores) Pteridophyta consist of two lineages (subphyla): microphyllous lycophytes and megaphyllous molinophytes

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 31 / 32 Plant diversity “Ferny” ferns For Further Reading

A. Shipunov. Introduction to Botany [Electronic resource]. 2010—onwards. Mode of access: http://ashipunov.info/shipunov/school/biol_154

Th. L. Rost, M. G. Barbour, C. R. Stocking, T. M. Murphy. Plant . 2nd edition. Thomson Brooks/Cole, 2006. Chapter 23.

Shipunov (MSU) Introduction to Botany. Lecture 29 November 14, 2014 32 / 32