The ANITA grade and the Magnoliid Complex

1. The ANITA grade and the Magnoliid Complex Spring 2011

2. Major Angiosperm Clades Amborellaceae Nymphaeales Austrobaileyales MAGNOLIID COMPLEX MONOCOTS EUDICOTS [TRICOLPATES] ANITA GRADE Soltis et al. 2000, APG II 2002, Judd et al. 2002

3. Basal Angiosperms:Nymphaeaceae(Water Lily Family) Widespread, tropics to temperate regions Aquatic rhizomatous herbs, sap milky 70 species; 8 genera Flowers: many parts; laminar stamens; “floating”; colorful perianth; “beetle” pollination syndrome Special uses: ornamentals; sacred lotus Required taxa: Nymphaea (water lily)

4. 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)

5. Nymphaea odorata – Water Lily

6. 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 Table 4.1 in Judd et al.

7. Nymphaea tuberosaWater Lily

8. NupharWater Lily

9. Victoria amazonica – Giant Water Lily

10. Major Angiosperm Clades Amborellaceae Nymphaeales Austrobaileyales MAGNOLIID COMPLEX MONOCOTS EUDICOTS [TRICOLPATES] ANITA GRADE Soltis et al. 2000, APG II 2002, Judd et al. 2002

11. Angiosperm Evolution: Pollen • Basic division in distinguishing angiosperms is NOT monocotversusdicot!! • Basic distinction is the number of pores or sulcae (grooves) in the pollen grains. • Trends in pollen evolution clearly show a shift from uniaperturate pollen found in gymnosperms and the basal angiosperms to triaperturate pollen found in the more derived flowering plants. • Plesiomorphic condition in angiosperms is one sulcus (monocolpate). • Fossil record is quite good to document this transformation in pollen type.

12. Figure 9.1 from the text, Showing monocolpate to Tricolpate pollen

13. 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)

14. 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 genera, 220 species • Flowers: apocarpous; anthers laminar, large number; receptacle elongated • Significant features: Aromatic; fruit an aggregate of follicles in Magnolia or winged samaras in Liriodendron • Special uses: ornamentals; timber • Required taxa: Magnolia

15. Magnoliaceae • solitary flower • elongate receptacle • aggregate of follicles • woody plant • simple leaves • stipules • many spirally arranged parts • separate carpels • laminar stamens Magnolia virginiana sweetbay

16. Magnolia X soulangeana

17. Within the family, Magnolia is diagnosed by: -presence of a red or orange fleshy seed coat -follicles opening along the abaxial seam

18. Liriodendron tulipiferaTulip Poplar Tree

19. Magnoliids—Piperales:Piperaceae • Widely distributed in tropical and subtropical regions. • Primarily herbs; sometimes epiphytic; small trees • Diversity: 2,020 species, 6 genera • Flowers: minute, densely packed in thick spikes; noperianth (!); 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; for information only

20. Peperomia

21. Piper nigrum – Black pepper

22. 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 more or less inferior, of (4-)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. • Required taxa: Asarum, Aristolochia

23. Asarum (wild ginger) -stemless perennial with aromatic rhizomes -flowers actinomorphic -filament tips extending beyond the anthers -ovary inferior -fruit a fleshy capsule, seeds large

24. Characters of Aristolochia(Dutchman’s pipe) • Tropical or warm temperate regions • Perennial herbs or shrubs, twining or climbing or sometimes upright • Calyx tubular, greenish or purplish • Anthers sessile, strongly adnate to the short and fleshy style • “trap” flowers

25. Ceratophyllaceae -Submerged aquatic with many adaptations for this habitat -Fossil record extends back to the early Cretaceous -Phylogenetic position uncertain, but clearly part of the early radiation of angiosperms above the ANITA grade

26. On to the monocots… ...clearly monophyletic…superbly apomorphic! ! !