The Angiosperms Part II The Angiosperms Some terminology… Angiosperm phylogeny (part II) Evolutionary trends (part II) The Angiosperms Some terminology… Angiosperm phylogeny (part II) Evolutionary trends (part II) epiphyte -grows entirely in tree, with no connection to ground hemiepiphyte -starts in trees, sends down roots -this part is a root Angiosperm growth forms liana -woody climber vine -herbaceous climber tree shrub -a single trunk many stems liana -starts on ground, grows up tree -this part is a stem Compound vs. simple leaves blade blade petiole petiole Simple leaf Compound leaf A few more floral terms Hypanthium & bract petals sepals hypanthium bract Placentation placenta Placenta • tissue which bears the ovules Placentation • arrangement of placenta and ovules inside the ovary Ovary position Floral symmetry Radial • many planes of symmetry superior Bilateral • only one plane of symmetry inferior The Angiosperms Some terminology… Angiosperm phylogeny (part II) Evolutionary trends (part II) Monocots vs. “dicots” • • • • • • • flower parts in 3s carpels fused, 3 1 cotyledon (first leaf) parallel leaf veins mostly herbs scattered vascular bundles lack true secondary growth cotyledon most other angiosperms ba eu sal dic ots Mo no co ts Ma gn oli ids AN IT A gra de “Dicots” Rosids Asterids Remember: Monocots are monophyletic, “dicots” are not yll ale s Ca ryo ph ba eu sal dic ots Mo no co ts Ma gn oli ids AN IT A gra de Eudicots Rosids Asterids The Eudicots • triaperturate pollen • Petals and sepals in multiples of 4 and 5 Mo no co ts Ma gn oli ids Am bo rel la Ny mp ha ea Illi ce ciu ae m Eudicots triaperturate pollen uniaperturate pollen gra de Ca ryo ph ba eu sal dic ots yll ale s Mo no co ts Ma gn oli ids AN IT A Eudicots Rosids Asterids The Basal Eudicots The Basal Eudicots A mix of primitive and derived features Primitive: • Carpels numerous & separate • Stamens numerous, spiralled Advanced • filaments more or less defined • petals reduced in number • • Many exceptions to the above characters! Primitive characters lost in the advanced dicots still persist within the basal dicots Example Basal Eudicots Ranunculaceae (Buttercups) Papaveraceae (Poppy family) Platanaceae (Sycamore) gra de Ca ryo ph ba eu sal dic ots yll ale s Mo no co ts Ma gn oli ids AN IT A Eudicots Rosids Asterids The Caryophyllales Caryophyllales • Carpels fused • Unusual free-central placentation • Betalain pigments (unique to this order) • Many (most?): no petals! – Petal-like parts of flowers derived from sepals, stamens, or floral bracts – Can be free or tubular • Many species use CAM and C4 photosynthesis Caryophyllaceae free-central placentation “petals” derived from sterile stamens Nyctaginaceae petal-like structures derived from floral bracts and sepals bract sepals Red Pigments Betalains all Caryophyllales except Caryophyllaceae Anthocyanins All other angiosperms + Caryophyllaceae independent evolution? Caryophyllales Nyctaginaceae Cactaceae Caryophyllaceae gra de Ca ryo ph ba eu sal dic ots yll ale s Mo no co ts Ma gn oli ids AN IT A Eudicots Rosids Asterids The Rosids Rosids • • • • • • Huge, morphologically-diverse clade Carpels (separate but usually few) to fully fused Stamen few to many, but filament generally well-defined Petals free (mostly) Compound leaves common Hypanthium usually present hypanthium Hypanthium Pyrus (Rosaceae) Myrtales • Herbs, shrubs or trees • Inferior ovary + hypanthium • Hypanthium typically functions in pollination as nectar container or spur Myrtaceae Melastomataceae Onagraceae Floral tube is formed from hypanthium, not from corolla Longitudinal section of Fuchsia flower (Onagraceae) Fabales: Fabaceae • Bean family • “Legume” fruit • Herbs to rainforest canopy trees • 18,000 species • The dominant family (most individuals and species) in most tropical & subtropical forests • N-fixation common Fabaceae (pea family) Compound leaves Legume fruit calyx Subfamily Mimosoideae •one of few examples of truly tubular corolla outside the Asterids (convergent!) corolla Fagales • Trees and shrubs • Wind pollinated • Flowers tiny, in catkins • N-fixers • Commonly symbiotic with root fungi: assist uptake of soil nutrients Alder (Alnus) Oaks, walnuts, alder, beech, poplar Oak (Quercus) Fagales are dominant in many forests of temperate zone and in some tropical mountains Oak forest, Costa Rica Brassicales • Most are herbs • Mustard oils: unique to order • Includes mustard, broccolli, cauliflower, cabbage, and papaya Brassicaceae Caricaceae (papaya) Brassicaceae Mustard oils (Brassicales) Characteristic odor and taste of mustard, horseradish, cabbage Ca ryo ph ba eu sal dic ots yll ale s Mo no co ts AN IT A gra Ch de lor an tha Ma les gn oli ids Eudicots Rosids Asterids The Asterids Asterids • Tubular corolla • Stamens borne on petals • # stamens = or < # petals (except Ericales) • Style usually single • Carpels commonly reduced to 2 • Iridoid compounds Irididoid compounds • • Restricted to the asterids many different varieties, especially in Asteraceae Ericales • “Basal” asterids Theaceae • Some have separate petals • Stamens frequently > number of petals Ericaceae (Blueberry family) • note single style • anthers open by pores Fouqueriaceae (ocotillo) Gentianales • 14,000 species • Mostly tropical Apocynaceae • Simple opposite leaves with stipules Gentianaceae Rubiaceae Stipule Rubiaceae Solanales • Radially symmetrical flowers with folds in corolla • Mostly simple alternate leaves without stipules • Rich in alkaloids Convolvulaceae Solanaceae: chile Solanaceae: tomato Solanaceae Alkaloids • Toxic defensive compounds • Many are psychoactive (hallucinogenic) • Common in Solanaceae, but have evolved many times in different eudicot lineages Solanaceae (asterid) Cactaceae (basal eudicot) Lamiales • Herbs to lianas to huge trees • Bilaterally symmetrical flowers • blah Lamiaceae (Mint family) Bignoniaceae Bignoniaceae Acanthaceae Asteraceae (sunflowers) • • • Largest or second-largest plant family (20-25,000 species) worldwide distribution, temperate zone and tropics most are herbs, but also some weak-wooded trees Asteraceae (sunflowers) • • flowers in “heads” can contain 2 types of flowers: disc florets ray florets Asteraceae disk floret Asteraceae are dominant in many open landscapes (prairie, desert, tropical alpine) The Angiosperms Some terminology… Angiosperm phylogeny (part II) Evolutionary trends (part II) Evolution of the carpel leaf-like, separate enclosed, with fusion of multiple carpels Evolution of the stamen leaf-like clearly differentiated into specialized filament and anthers Evolution of the Angiosperm flower Floral reduction and fusion • Carpels: many, separate to few, fused • Perianth parts: all alike, to clearly differentiated sepals and petals • Petals: many, free to few, fused • Ovary: superior to inferior Eudicots ba eu sal dic ots Mo no co ts Ma gn oli ids AN IT A gra de Evolution of the angiosperm flower: carpel Rosids Asterids ovary with 2 carpels carpel fusion and reduction carpel fusion and reduction carpels separate, usually numerous Eudicots ba eu sal dic ots Mo no co ts Ma gn oli ids AN IT A gra de Evolution of the angiosperm flower: stamens Rosids stamens with distinct, threadlike filament stamens with distinct, thread-like filament stamens laminar, undifferentiated Asterids Eudicots ba eu sal dic ots Mo no co ts Ma gn oli ids AN IT A gra de Evolution of the angiosperm flower: perianth Rosids Asterids fused corolla distinct sepals and petals flower parts in 4s & 5s flower parts in whorls of 3 perianth undifferentiated, spiralled, numerous Convergent evolution: succulent growth form Euphorbia • stem succulent Senecio (Asteraceae) • stem and leaf succulents Cactaceae • stem succulent Aloe •leaf succulent Convergent evolution: petal-like structures Nyctaginaceae •petaloid floral bracts •petaloid sepals Caryophyllaceae •sterile stamens (staminodia) Zingiberaceae •sterile stamens (staminodia) Convergent evolution: the fused tubular flower Onagraceae •tubular extension of apex of ovary wall (hypanthium) Solanaceae •fused petals Nyctaginaceae •fused sepals •(plus sepal-like bracts) Amaryllidaceae •fused sepals and petals Questions 1. What characters distinguish the basal eudicots, the Caryophyllales, the Rosids, and the Asterids 2. What floral characters distinguish a “primitive” ANITA angiosperm such as Illicium or the water lilies from an “advanced” angiosperm such as an Asterid? 3. Provide examples of convergent evolution among the Eudicots, and examples of taxa where they occur