The ANITA grade and the Magnoliid Complex Spring 2013 Major Angiosperm Clades Amborellaceae Nymphaeales ANITA GRADE Austrobaileyales MAGNOLIID COMPLEX MONOCOTS Soltis et al. 2000, APG II 2002, Judd et al. 2002 EUDICOTS [TRICOLPATES] Angiosperm Evolution: Pollen • • • • • Basic division in distinguishing angiosperms is NOT monocot versus dicot!! Basic distinction is the number of pores or sulcae (grooves) in the pollen grains. Trends in pollen evolution clearly show a shift from uni-aperturate pollen found in gymnosperms and the “basal” angiosperms and monocots to tri-aperturate pollen found in the eudicots. Plesiomorphic condition in angiosperms is uniaperturate. Fossil record is quite good to document this transformation in pollen type. Fig. 7.1 monoaperturate pollen (one pore or groove) Pollen Types Fig. 7.1 “Basal” Angiosperms: Amborellaceae (Amborella Family) Amborella trichopoda Fig. 7.1 “Basal” Angiosperms: Nymphaeaceae (Water Lily Family) • Widespread, tropics to temperate regions • Aquatic rhizomatous herbs, sap milky • Ca. 70 species; 5-6 genera • Flowers: many parts; laminar stamens; • • floating; colorful perianth; beetle pollination syndrome Special uses: ornamentals Required family “Basal” Angiosperms: Nymphaeaceae (Water Lily Family) • • • • • • • Widespread, tropics to temperate regions Aquatic herbs with floating leaves, often with milky sap (latex) Number of species: ca. 70 species; 5-6 genera Flowers: floating; many parts; colorful perianth laminar stamens; gynoecium syncarpous with 3many carpels Significant features: beetle pollination syndrome Special uses: ornamentals, edible rhizomes and seeds Required family “Basal” Angiosperms: Nymphaeaceae (Water-lily Family) •numerous petals, stamens, carpels •laminar stamens •pollen monosulcate •stigma discoid, radiating •berry-like fruit, dehiscent •perisperm (from the sporophyte) •usually lack vessels (or have tracheidlike vessels) Nymphaea odorata – Water Lily Beetle pollination syndrome • Flowers often green or white • Flowers with various but strong scents • Can open during the day or night • Flowers open or enclosed • Nectar and/or pollen reward • See Ch. 13 in Simpson Beetle pollination in water lilies: http://vimeo.com/41976231 Double flowering to avoid selfing: http://www.youtube.com/watch?v=B2S5c1s5mPA Nymphaea tuberosa Water Lily Nuphar Water Lily Victoria amazonica – Giant Water Lily Major Angiosperm Clades Amborellaceae Nymphaeales ANITA GRADE Austrobaileyales MAGNOLIID COMPLEX MONOCOTS Soltis et al. 2000, APG II 2002, Judd et al. 2002 EUDICOTS [TRICOLPATES] Fig. 7.1 Magnoliid characters “Magnocots” •2-ranked leaves, paracytic stomates •Perianth generally 3-merous •Stamens and carpels: distinct, numerous, spirally arranged •Boat-shaped, monosulcate pollen •Superior ovary •Seeds with fleshy seed coat/aril in many; minute embryo, copious endosperm •Many anatomical characters (esp. wood) Magnoliids—Magnoliales: Magnoliaceae (The Magnolia Family) • Temperate to tropical regions of eastern North America; east Asia, South America • Trees or shrubs with simple leaves • Number of species: 2 or 7 genera, 200 species • Flowers: apocarpous; anthers laminar, many; receptacle elongated • Significant features: Aromatic; fruit an • • aggregate of follicles in Magnolia or winged samaras in Liriodendron Special uses: ornamentals; timber Required family Magnoliaceae •woody plant •solitary flower •simple leaves •stipules •many spirally arranged parts Magnolia virginiana sweetbay •elongate receptacle •separate carpels •laminar stamens •aggregate of follicles Magnolia X soulangeana Magnolia Dispersal mechanism? Liriodendron tulipifera Tulip Poplar Tree Other Magnoliid families of interest (but not required) • Lauraceae (the laurel or bay family) • Piperaceae (the pepper family) • Aristolochiaceae (the birthwort family) Magnoliids—Laurales: Lauraceae (Laurel or Bay Family) • • • • • • • Widespread in tropical and subtropical regions; SE Asia & northern South America Trees, shrubs, vines. Diversity: 2,200 species, 45 genera Flowers: concave receptacle that often enlarges in fruit; anthers dehiscing via pores with flaps; sticky pollen; pollen without apertures; carpel 1; embryo large, endosperm lacking. Significant features: ethereal oils Special uses: cinnamon (Cinnamomum) and bay (Laurus nobilis) used as spices; avocado (Persea) Family not required Lauraceae Sassafras albidum -original source of root beer flavor -ground leaves→ filé powder Lauraceae Economic plants and products: Cinnamomum zeylandicum True cinnamon Bird dispersal and ecological role of Lauraceae Magnoliids—Piperales: Piperaceae • • • • • • • Widely distributed in tropical and subtropical regions. Primarily herbs; sometimes epiphytic; small trees Diversity: 3,600 species, 5 genera Flowers: minute, densely packed in a thick spike or spadix; no perianth (!); carpels 1-4, connate, with 1 ovule per gynoecium, basal placentation; fruit usually a drupe. Significant features: stem with bundles in >1 ring or + scattered Special uses: Piper nigrum (black pepper); Piper betle (betel pepper), ornamentals (Peperomia) Family not required Piperaceae - Peperomia Peperomia leaves Piper nigrum – Black pepper Magnoliids—Piperales: Aristolochiaceae • • • • • • • (Dutchman’s Pipe or Birthwort Family) Widespread in tropical and subtropical regions; absent in Australia. Lianas or herbs, occasionally shrubs Diversity: ca. 460 species, 7 genera Flowers: Highly modified, showy, fused sepals, radial or bilateral, tubular, and S-shaped or pipe shaped calyx tube; corolla usually lacking or vestigial; ovary inferior or half-inferior, of 3-6 connate carpels; ovules numerous; filaments more or less adnate to style. Significant features: “dead meat” carrion coloration attracts insects, usually flies. Trap-flowers. Special uses: ornamentals; some medicinal uses. Family not required Asarum (wild ginger) -perennial with aromatic rhizomes -flowers actinomorphic -ovary inferior -fruit a fleshy capsule, seeds large & with elaiosomes, ant-dispersed Aristolochia (Dutchman’s pipe) Trap flowers On to the monocots… ...clearly monophyletic…superbly apomorphic! ! !