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The Monocots: Part 2 Commelinoid Monocots
Spring 2010
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Phylogeny of Monocot Groups
Acorales Alismatales Asparagales Liliales Dioscoreales Pandanales Arecales Poales Commelinales Zingiberales Basal “Petaloid” Commelinoid
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Commelinoid characters
Special type of epicuticular wax Starchy pollen UV-fluorescent compounds in the cell walls Starchy endosperm (except in the palms) Lots of molecular support
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Commelinoid Monocot Groups
Order Arecales - Palms Arecaceae (Palmae) Order Poales - Grasses - Bromeliads Cat-tails Rushes, Sedges, and Grasses Bromeliaceae Typhaceae Juncaceae Cyperaceae Poaceae (Gramineae) Order Zingiberales – Ginger, banana, and allies (no required families)
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Commelinoid Monocots: Arecales: Arecaeae (Palmae)
Widespread throughout tropical and warm temperate regions “Trees” or “shrubs”, typically unbranched Diversity: ca. 2,780 in 200 genera Flowers: usually sessile, in compound-spicate inflorescences, these subtended by a bract (spathe); ovule 1 per locule Significant features: Leaves alternate or spiral, blades plicate, splitting in a pinnate or palmate manner Special uses: coconut (Cocos nucifera), date (Phoenix dactylifera), rattan (Calamus), oils and waxes, ornamentals Required taxa: *family only * Change from lab manual
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Arecaceae Numerous small flowers
Spathes + compound-spicate inflorescence 3 sepals + 3 petals Superior ovary (carpel fusion varies) Drupe Unbranched trunks Big leaves on top!
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Arecaceae – The Palm Family
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Arecaceae – Cocos nucifera
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Arecaceae Economic plants and products: Phoenix dactylifera Dates
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Characters of Poales Silica bodies (in silica cells) in the epidermis
Styles strongly branched Loss of raphide (needle-like) crystals Much molecular support for monophyly Wind pollination has evolved several times independently within the order Ecologically very important
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Commelinoid Monocots—Poales: Bromeliaceae (The Pineapple/Bromeliad Family)
Tropical to temperate regions of the Americas Predominantly epiphytic herbs (“tank” plants) Diversity: ca. 1,520 species in 51 genera Flowers: radial, perianth differentiated into calyx and corolla, borne in axils of often brightly colored bracts; inflorescences spicate or paniculate; stigmas 3, usually twisted; seeds often winged or with tufts of hair Significant features: leaves with water absorbing peltate (or stellate) scales Special uses: pineapple (Ananas) Required taxa: *Tillandsia (Spanish moss) *change from lab manual
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Bromeliaceae
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Bromeliaceae: Tillandsia (Spanish moss)
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Bromeliaceae – Ananas comosus
Fruit type?
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Commelinoid Monocots—Poales: Typhaceae (The Cattail Family)
Widely distributed, especially in Northern Hemisphere Aquatic & wetland rhizomatous herbs Diversity: 28 species in 2 genera Flowers: small, unisexual; separated spatially on dense, compact spicate or globose-clustered inflorescences; placentation apical Significant features: rhizomatous; long slender leaves; characteristic inflorescence Special uses: ornamental aquatics Required taxa: Typha
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Typhaceae - Typha
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Commelinoid Monocots—Poales: Juncaceae (The Rush Family)
Worldwide, mostly temperate regions; wet or damp habitats Rhizomatous herbs, stems round and solid Diversity: 400 species in 6 genera Flowers: tepals 6, distinct; carpels 3 in superior ovary; stamens 6; fruit a loculicidal capsule Significant features: leaves 3-ranked, sheaths usually open Special uses: leaves used to weave rush baskets; some ornamentals Required taxa: Juncus
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Juncaceae
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Juncaceae: Juncus -cymose inflorescences -leaf sheaths open
-leaf blades flat, grooved, or cylindrical
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Commelinoid Monocots—Poales: Cyperaceae (The Sedge Family)
Worldwide, usually in damp or semi-aquatic sites Rhizomatous herbs, stems usually triangular in cross section Diversity: 4,500 species in 104 genera Flowers: with subtending bract; tepals absent or reduced to 3-6 scales or hairs; stamens 1-3; carpels 2-3 in superior ovary; fruit an achene (nutlet) Significant features: Inflorescence a complex group of spikelets; leaf sheaths closed, ligule lacking; silica bodies conical Special uses: Papyrus used originally for paper; “water chestnuts”and a few other rhizomes edible, leaves used for weaving; some ornamentals. Required taxa: Carex, Cyperus
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Cyperaceae versus Juncaceae: Field Character
“Sedges have edges… …and rushes roll.”
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Cyperaceae
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Cyperaceae Flowers: Arranged in spikelets Reduced
Wind-pollinated flowers Subtended by bract Reduced/absent perianth flower + subtending bract = floret flower spikelet
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Cyperaceae Fruit type is the achene: very important in
the taxonomy of the family. Eleocharis Rhynchospora (note bristle perianth) Cyperus
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Cyperaceae: Cyperus
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Cyperaceae: Carex -presence of the perigynium (a sac-like
bract surrounding the female flower) in addition to the subtending bract -leaves usually with a ligule
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Commelinoid Monocots—Poales: Poaceae (Gramineae) (The Grass Family)
Cosmopolitan Primarily herbs, often rhizomatous; “trees” in most bamboos; stems are called culms Diversity: 10,000 species in ca. 650 genera Flowers: small, perianth parts reduced to lodicules; each flower enclosed by two bracts (lemma and palea) = floret; stamens typically 3; carpels 3, but appearing as 2; fruit a caryopsis Significant features: 1-many florets aggregated into spikelets, each with usually 2 empty bracts (glumes) at the base; leaf with a ligule Special uses: many – grains, turf, fodder/forage, structural uses (e.g., bamboo). Required taxa: *Poa, *Andropogon *change from lab manual
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Economic importance bamboo sugar cane Zea mays weeds Oryza sativa
Triticum aestivum
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Ecological importance
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Poaceae: vegetative structure
ligule
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Poaceae: spikelet and flower structure
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Anatomy of the Caryopsis (Grain)
The fruit wall (pericarp) is completely fused to the seed coat. Endosperm (3N; triploid) contains the bulk of starch storage in the seed. The embryo is a pre-formed grass plant, with apical meristems (for both shoot and root) and protective organs (coleoptile and coleorhiza) which emerge first during germination.
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Origin of grasses ca. 70-80 mya in southern- hemisphere forests
early grasses Origin of grasses ca mya in southern- hemisphere forests
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Origin of grasses ca. 70-80 mya in forests Anomochlooideae Pharoideae
Puelioideae Bamboos (Bambusoideae) Origin of grasses ca mya in forests Bluegrasses (Pooideae) Rices (Ehrhartoideae) Panicgrasses (Panicoideae) Major radiation in Oligocene- Miocene epochs into open habitats Needlegrasses (Aristidoideae) Lovegrasses (Chloridoideae) + Micrairoideae Stamens reduced to 3 Reeds (Arundinoideae) Oatgrasses (Danthonioideae)
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C4 photosynthetic pathway
(in warm season grasses) is advantageous under higher temperatures, higher light, and less water
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Dispersal!
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Poaceae (Gramineae)
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Poaceae: Poa -cool season -leaf tip boat-shaped
-inflorescence branched from a main axis -spikelets solitary -glumes papery -florets 3-several, often with a cottony web at the base
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Poaceae: Andropogon -warm season -leaf midrib whitish, prominent
-2-many branches per inflorescence, often digitate -spikelets paired -glumes tough, leathery
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For more information and images: http://www. eeob. iastate
For more information and images: The Grasses of Iowa
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Grasses, Sedge, Rushes! Stem terete, hollow, or solid, jointed
Leaf ranks 2 Leaf sheath Open, ligule Inflor: Spikelets Perianth: Lodicules Fruit: Caryopsis Triangular, solid, not jointed 3 Closed Spikelets None or bristles/scales Achene Terete, solid, not jointed 3 Open Cymose 6 chaffy tepals Capsule
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“Graminoids” - Comparison
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Commelinales 5 families, 780 species, widespread in various habitats
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Commelinoid Monocots: Zingiberales
Large herbs with vessels more or less limited to the roots Silica cells present in the bundle sheaths Leaves clearly differentiated into a petiole and blade Leaf blade with pinnate venation, often tearing between the second-order veins Leaf blade rolled into a tube in bud Petiole with enlarged air canals Flowers bilateral (or irregular) Pollen lacking an exine Ovary inferior Seeds arillate and with perisperm 8 families and nearly 2000 species Must be able to recognize the order!
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Zingiberales diversity
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Musaceae Musa
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