The ANITA grade and the Magnoliid Complex

Spring 2012

Major Angiosperm Clades

Amborellaceae Nymphaeales Austrobaileyales ANITA GRADE

Soltis et al. 2000, APG II 2002, Judd et al. 2002

MAGNOLIID COMPLEX MONOCOTS 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 to tria-perturate pollen found in the more derived flowering plants.

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 taxa: Nymphaea (water lily)

“Basal” Angiosperms: Nymphaeaceae (Water-lily Family)

•

numerous petals, stamens, carpels

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laminar stamens

•

pollen monosulcate

•

stigma discoid, radiating

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berry-like fruit, dehiscent

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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

Nymphaea tuberosa

Water Lily

Nuphar

Water Lily

Victoria amazonica

– Giant Water Lily

Major Angiosperm Clades

Amborellaceae Nymphaeales Austrobaileyales ANITA GRADE

Soltis et al. 2000, APG II 2002, Judd et al. 2002

MAGNOLIID COMPLEX MONOCOTS EUDICOTS [TRICOLPATES]

Fig. 7.1

Magnoliid characters “Magnocots”

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2-ranked leaves, paracytic stomates

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Perianth generally 3-merous

•

Stamens and carpels: distinct, numerous, spirally arranged

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Boat-shaped, monosulcate pollen

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Superior ovary

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Seeds with fleshy seed coat/aril in many; minute embryo, copious endosperm

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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, 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

•

woody plant

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simple leaves

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stipules

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many spirally arranged parts

•

separate carpels

•

laminar stamens

Magnoliaceae

•

solitary flower

Magnolia virginiana

sweetbay •

elongate receptacle

•

aggregate of follicles

Magnolia X soulangeana

Magnolia

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

Other Magnoliid families of interest (but not required)

•

Lauraceae (the laurel or bay family)

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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

Sassafras albidum

Cinnamomum

cinnamon

Lauraceae

Enlarged receptacle under the drupe!

Sassafras albidum

sassafras

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

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)

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

Characters of Aristolochia

• • • • •

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

On to the monocots…

...clearly monophyletic…superbly apomorphic! ! !