The Wonderful World of Green Plant Diversity and Evolution Biol 366 Spring 2012 Tree of Life: The Big Picture now Bacteria Archaea Eukaryotes >2 bya >3.5 bya ca. 4 bya membrane-bound nucleus, organelles, etc. Fig. 1.1 from Simpson Green plants share: • Chlorophylls a (ancestral) and b • Starch storage • Stellate flagellar structure • Certain gene transfers from the chloroplast to the nucleus • And other features (see Ch. 3) Green plant diversity: • Ca. 350,000 species • Two major groups: 1) chlorophytes (marine and other green algae) and 2) streptophytes [freshwater green algae and embryophytes (= land plants)] • A major branch (clade) in the eukaryotic Tree of Life Fig. 3.1 from Simpson Some definitions • Clade = branch on an evolutionary tree, a lineage, includes an ancestor and all its descendants. Ex.: Green plants, chlorophytes, land plants. • Paraphyletic group = a group that includes an ancestor and some (but not all) of its descendants, indicated by double quotation marks. Ex.: “Green algae” Chlorophytes Fig. 3.1 from Simpson Spirogyra Chara Nitella (Judd et al. 2008) Basal streptophytes desmids haploid body Conjugation in Spirogyra Haplontic life cycle (haploid dominant or zygotic meiosis) The only diploid cell Is the zygote zygote (2n) biology.unm.edu ♀ Charales Haplontic but some have multicellular gametangia (gamete-producing structures) ♂ mason.gmu.edu Embryophytes (land plants) share: • Cuticle • Alternation of generations (multicellular sporophyte and multicellular gametophyte) • Multicellular gametangia (gamete-producing structures) • Multicellular sporangium (spore-producing structure) • Embryo (young sporophyte) • Parenchyma? (more likely ancestral) Generalized embryophyte life cycle: Alternation of generations Bryophytes • Hornworts, liverworts, mosses • Gametophyte-dominant • No vascular tissue (except conducting cells in a few mosses) • Separate male and female gametophytes • Sperm must swim to the egg, therefore need water for fertilization and therefore must remain small liverworts mosses hornworts Plant Tree of Life: Embryophtes “Bryophytes” now Liverworts ca. 450 mya Hornworts Mosses Tracheophytes (vascular plants) Liverwort gametophyte Liverwort thallus showing air pores Liverwort Multicellular gametangia (male = antheridia) Liverwort Multicellular gametangia (female = archegonia) Oogamy Retention of zygote within the female gametophyte Multicellular embryo G S Hornworts Moss male gametangia (= antheridia) Capsule = sporangium of the sporophyte Generalized embryophyte life cycle: Alternation of generations Tracheophytes (vascular plants) • Vascular tissue (tracheids) present • Include lycophytes (quillworts, clubmosses, spikemosses), monilophytes (ferns, horsetails, whisk ferns), and spermatophytes (seed plants) Fig. 4.1 from Simpson Lycophytes & Monilophytes • Quillworts, clubmosses & spikemosses (= lycophytes); ferns, whisk-ferns, & horsetails (= monilophytes); • Independent gametophytes and sporophytes • Sperm must still swim to the egg • Most are homosporous; a few evolved heterospory • Many homosporous ferns have means of avoiding self-fertilization Lycophytes Isoetes (quillwort) Lycopodium and friends (clubmosses) Selaginella (spikemoss) Monilophytes (ferns, horsetails, whisk ferns) horsetails Whisk-fern (Psilotum) Ferns (Leptosporangia) Nutritionally independent sporophytes and 2n gametophytes sporophyte 2n gametophyte 1n 1n spores 2n 2n Fern Life Cycle, Fig. 4.32, Simpson Lignophytes (woody plants) & Spermatophytes (seed plants) • Secondary xylem (wood) & bark, heterospory, seeds, eustele, pollen (also pollen tube, pollination droplet) • Includes gymnosperms and angiosperms Fig. 5.1 from Simpson Gymnosperms • Conifers, gingko, cycads, Gnetales • Molecular data support this group as having a single common ancestor • No obvious defining character (see characters for Lignophytes & Spermatophytes) Female cone with each scale bearing usually two ovules; directly exposed to pollen Male cones with each scale bearing two or more microsporangia male pine pollen pine microsporangia biology.ualberta.edu female Fig. 5.7 from Simpson Angiosperms • • • • • “Dicotyledons”, monocotyledons Heterosporous (ancestral) Sporophyte-dominant (ancestral) Pollen = male gametophyte (ancestral) Archegonia lost; embryo sac = female gametophyte; ovules enclosed in carpels (indirect pollination) • Double fertilization produces zygote + primary endosperm nucleus Flower = a short, determinate shoot bearing highly modified leaves, some of which are fertile (i.e., bearing either microsporangia or megasporangia), with the megasporangia in carpels Animal pollination syndromes Wind pollination A wide range of fruit types… Fig. 5.7 from Simpson The wonderful world of land plant diversity over 300,000 species of angiosperms