The ANITA grade and the Magnoliid Complex Spring 2014
Major Angiosperm Clades Amborellaceae Nymphaeales Austrobaileyales MAGNOLIID COMPLEX MONOCOTS EUDICOTS [TRICOLPATES] ANITA GRADE Soltis et al. 2000, APG II 2002, Judd et al. 2002
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 uni-aperturate. 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 3-many 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 tracheid-like 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
http://vimeo.com/41976231 Beetle pollination in water lilies: 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 Austrobaileyales MAGNOLIID COMPLEX MONOCOTS EUDICOTS [TRICOLPATES] ANITA GRADE Soltis et al. 2000, APG II 2002, Judd et al. 2002
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
aggregate of follicles many spirally arranged parts Magnoliaceae solitary flower elongate receptacle aggregate of follicles woody plant simple leaves stipules many spirally arranged parts separate carpels laminar stamens Magnolia virginiana sweetbay
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
Economic plants and products: Cinnamomum zeylandicum 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! ! !