The ANITA grade and the Magnoliid Complex Spring 2011

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

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)

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 Table 4.1 in Judd et al.

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 10

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 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.

Figure 9.1 from the text, Showing monocolpate to Tricolpate pollen

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 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

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

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

Liriodendron tulipifera Tulip Poplar Tree

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; 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; for information only

Peperomia

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 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

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

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

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

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