Diversity And Classification of Flowering Plants: Amborellales, Nymphaeales, Austrobaileyales, Magnoliids, Ceratophyllales Michael G. Simpson.
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Diversity And Classification of Flowering Plants: Amborellales, Nymphaeales, Austrobaileyales, Magnoliids, Ceratophyllales Michael G. Simpson How do we know angiosperm relationships? Cladistic analyses: parsimony, maximum likelihood, Bayesian Use of all types of data: morphological, anatomical, embryological, palynological, karyological, chemical, and molecular data Most useful: multiple gene sequence data Angiosperm Phylogeny Group (APG III) system - 2009 Use of orders (end in “-ales”) to classify one or more families Orders are well-supported based on measures of clade robustness (bootstrap, jackknife) Some orders further grouped into higher taxa (e.g., monocots, Asterids, etc.) Floral Formulas Measures number of parts (merosity & cycly): K = calyx C = corolla P = perianth A = androecium G = gynoecium ( ) = fusion of like parts [ ] = less common E.g., • K (5) [(4)] C 5 [4] A 5+5 [4+4] G 5 [4], superior • P (3+3) A 3+3 G (3), inferior Floral Diagrams Cross-sectional representation of a flower Amborellales - the most basal, extant angiosperm lineage • Monotypic (1 family, 1 species) Amborellaceae Amborella trichopoda Native to New Caledonia Amborellaceae • The Amborellaceae are distinctive in being vesselless, evergreen shrubs with unisexual flowers having an undifferentiated, spiral perianth, numerous, laminar stamens, and an apocarpous, apically-open gynoecium, with 1-ovuled carpels. • Male flowers: P 5-8 A ∞ • Female flowers: P 5-8 G 5-6, superior Amborella trichopoda Amborella trichopoda Photo by Tim Stevens U. C. S. C. Photo by Missouri Botanical Garden Flowers unisexual: male (left) female (above) Amborella, if it represents a “primitive” angiosperm: • vessel-less (no vessels, only tracheids) • woody shrub with simple, glabrous, evergreen leaves • flowers unisexual • perianth undifferentiated • stamens numerous, laminar • gynoecium apocarpous • fruit an aggregate of drupes • seeds endospermous NYMPHAEALES Three families: Cabombaceae Hydatellaceae Nymphaeaceae Nymphaeaceae - Water-Lily family (Nymphe, a water nymph). 6 genera / 60 species The Nymphaeaceae are distinguished from related families in consisting of aquatic herbs with floating leaves and solitary, floating to emergent flowers with mostly spiral floral parts and petals grading into usually laminar stamens. K 4-6 [-14] C 8-∞ [0] A ∞ G (3-∞), superior or inferior. Nuphar luteum Spadder-dock Nymphaea spp. Water-lily Nymphaea sp. Nymphaea sp. Victoria reginae AUSTROBAILEYALES Austrobaileyaceae Illiciaceae Schisandraceae Trimeniaceae Illiciaceae The Illiciaceae are distinctive in being evergreen trees or shrubs with aromatic oil cells, glabrous, spiral pellucid-punctate, exstipulate leaves, and small flowers with numerous, spiral tepals (outer sepal-like, inner petal-like), few to numerous stamens, and few-numerous, one-seeded, apocarpous pistils, the fruit a follicetum. P ∞ [7-33] A ∞ [4-50] hypanthium absent. G ∞ [5-21], superior, Illiciaceae Illiciaceae Illicium verum, star anise - spice LAURALES Atherospermataceae Calycanthaceae Gomortegaceae Hernandiaceae Lauraceae Monimiaceae Siparunaceae Lauraceae- Laurel family (L. laurus, laurel or bay). 45 genera / 2200 species The Lauraceae are distinguished from related families in consisting of perennial trees or shrubs (rarely vines) with aromatic oil glands, evergreen leaves, an undifferentiated perianth, valvular anther dehiscence, and a single, superior ovary having one ovule per carpel with apical placentation, seeds lacking endosperm. P 3+3 [6, 2+2, or 3+3+3] A 3-12+ G 1 [-(3)], superior, rarely inferior, hypanthium present Lauraceae Economically imporant members Cinnamomum cassia Cinnamomum zelanicum Cinnamomum camphora Laurus nobilis Persea americana cassia cinnamon camphor bay / laurel avocado Laurus nobilis Laurel Lauraceae valvular anther dehiscence Laurus nobilis Laurel Lauraceae • Persea americana Avocado Lauraceae • Persea americana Avocado Lauraceae • Cinnamomum camphora Camphor Lauraceae MAGNOLIALES Annonaceae Degeneriaceae Eupomatiaceae Himantandraceae Magnoliaceae Myristicaceae Magnoliaceae - Magnolia family (after Pierre Magnol of Monpelier, 1638-1715) 7 genera / 200 species The Magnoliaceae are distinguished from related families in consisting of trees and shrubs with stipulate leaves, solitary flowers, a usually undifferentiated petaloid perianth with numerous tepals, and numerous, spiral stamens and an apocarpous gynoecium of numerous, spiral pistils born on elongate receptacular axis (torus or androgynophore); the fruit is an aggregate of follicles, berries, or samaras, seeds usu. with a sarcotesta. P ∞ A ∞ G ∞ [2-∞], superior. Magnolia grandiflora Magnolia grandiflora Magnolia grandiflora Michelia doltsopa Michelia doltsopa PIPERALES Aristolochiaceae (incl. Lactoridaceae) Hydnoraceae Piperaceae Saururaceae Aristolochiaceae - Birthwort family (Gr. aristos, best + lochia, childbirth, from resemblance of a species of Aristolochia to the correct fetal position). 5-8 genera / 465-480 species Calyx enlarged, petaloid Corolla reduced to absent Stamens adnate to style, forming gynostemium Ovary inferior to half-inferior Locules generally six. K (3) C 0 [3] A 6-∞, usu. adnate to style G (4-6), inferior (half-inferior). Aristolochia elegans (left) & A. gigantea (right) Dutchman’s Pipe Aristolochiaceae Aristolochia macrophylla Dutchman’s Pipe Aristolochiaceae Aristolochia trilobata Dutchman’s Pipe Aristolochiaceae Aristolochia trilobata Dutchman’s Pipe Aristolochiaceae Aristolochia californica in fruit Dutchman’s Pipe Aristolochiaceae Asarum canadense Aristolochiaceae Hexastylis minor Aristolochiaceae Piperaceae - Pepper family (piper, Indian name for pepper). 14 genera / 1940 species The Piperaceae are distinctive in having a spadix with numerous, very small, unisexual or bisexual flowers lacking a perianth. Economic importance includes Piper nigrum, the source of black and white pepper; other species are used for flavoring, medicinal plants, euphoric plants (e.g., Piper methysticum, kava), and cultivated ornamentals, e.g., Peperomia spp. P 0 A 3+3 (1-10) G 1 or (G) 3,4, superior. Black Pepper: Piper nigrum • • • • • • Family: Piperaceae Vine Part used: Fruit - drupe Native to Malabar coast (India) World’s most important spice Black pepper – unripe drupe (green) fermented, dried, ground • White pepper – ripe drupe (red) – outer layer removed, inner dried, ground Black Pepper: Piper nigrum drupe (of spadix) Piper nigrum unripe, dried unripe drupes unripe, fermented ripe, exo-, mesocarp removed, dried ripe drupes Piper [Macropiper] excelsum Piperaceae Piper [Macropiper] excelsum Piperaceae Peperomia sp. Piperaceae Saururaceae - Lizard's-Tail family (Gr. saur, lizard + our, tail, in reference to the tail-shaped inflorescence of Saururus cernuus). 4 genera / 6 species The Saururaceae are distinctive in being perennial herbs with a bracteate spike or raceme and with flowers lacking a perianth, the ovary solitary, many-ovulate, the fruit a capsule. P 0 A 3, 3+3, or 4+4 G (3-5), superior. Anemopsis californica Yerba Mansa Saururaceae Saururus cernuus Lizard’s Tail Saururaceae CERATOPHYLLALES Ceratophyllaceae only The Ceratophyllaceae are distinguished from related families in consisting of aquatic herbs with whorled, dichotomously branched, serrulate leaves and solitary, unisexual flowers. P (8-12) A 5-27 G 1, superior. Ceratophyllum demersum