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The Monocots: Part 2 Commelinoid Monocots Spring 2011 Phylogeny of Monocot Groups Basal “Petaloid” Commelinoid Acorales Alismatales Asparagales Liliales Dioscoreales Pandanales Arecales Poales Commelinales Zingiberales 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 Commelinoid Monocot Groups Order Arecales - Palms Arecaceae (Palmae) Order Poales - Grasses - Bromeliads Cat-tails Rushes, Sedges, and Grasses Typhaceae Juncaceae Cyperaceae Poaceae (Gramineae) Order Zingiberales – Ginger, banana, and allies (no required families) 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 Arecaceae •Numerous small flowers •Spathes + compound-spicate inflorescence •3 sepals + 3 petals •Superior ovary (carpel fusion varies) •Drupe •Unbranched trunks •Big leaves on top! Arecaceae – The Palm Family Arecaceae – Cocos nucifera Arecaceae Economic plants and products: Phoenix dactylifera Dates 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 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) Family not required; for information only Bromeliaceae – Ananas comosus Fruit type? 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 Typhaceae - Typha 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 Juncaceae: Juncus -cymose inflorescences -leaf sheaths open -leaf blades flat, grooved, or cylindrical 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 1 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 Cyperaceae versus Juncaceae: Field Character “Sedges have edges… …and rushes roll.” Cyperaceae diversity Flowers: •Arranged in spikelets •Reduced •Wind-pollinated flowers •Subtended by bract •Reduced/absent perianth spikelet flower + subtending bract = floret Cyperaceae flower From Zomlefer 1994 Cyperaceae Fruit type is the achene: very important in the taxonomy of the family. Cyperus Eleocharis Rhynchospora (note bristle perianth) Cyperaceae http://waynesword.palomar.edu/termfl3.htm Cyperaceae: Cyperus -leaves usually basal -ligules absent -spikelet scales distichous, each subtending a flower -spikelets flattened or cylindrical -flowers bisexual -no perigynium 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 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: Zea, Triticum, Oryza bamboo sugar cane weeds Economic importance Zea mays Oryza sativa Triticum aestivum Ecological importance Poaceae: vegetative structure ligule Poaceae: spikelet and flower structure flower Images from Grasses of Iowa 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. early grasses Origin of grasses ca. 70-80 mya in southernhemisphere forests Anomochlooideae Pharoideae Puelioideae Origin of grasses ca. 70-80 mya in forests Bamboos (Bambusoideae) Bluegrasses (Pooideae) Rices (Ehrhartoideae) Panicgrasses Major radiation in OligoceneMiocene epochs into open habitats + (Panicoideae) Needlegrasses (Aristidoideae) Lovegrasses (Chloridoideae) Micrairoideae Stamens reduced to 3 Reeds (Arundinoideae) Oatgrasses (Danthonioideae) C4 photosynthetic pathway (in warm season grasses) is advantageous under higher temperatures, higher light, and less water Dispersal! Poaceae (Gramineae) diversity Oryza (rice) -aquatic or wetland herbs -one floret per spikelet -spikelets strongly flattened Triticum (wheat) -annuals -dense inflorescences -spikelets sessile, one per node -2-9 florets per spikelet Zea (maize or corn) -male and female spikelets usually on separate inflorescences -female inflorescences axillary, enclosed in 1 or more sheaths (husks), one sessile spikelet per node -male inflorescences terminal, with paired spikelets For more information and images: http://www.eeob.iastate.edu/research/iowagrasses/ The Grasses of Iowa Grasses, Sedge, Rushes! •Stem terete, hollow, or solid, jointed •Triangular, solid, not jointed •Terete, solid, not jointed •Leaf ranks 2 •3 •3 •Leaf sheath Open, ligule •Closed •Open •Inflor: Spikelets •Spikelets •Cymose •Perianth: Lodicules •None or bristles/scales Achene •6 chaffy tepals •Fruit: Caryopsis •Capsule “Graminoids” - Comparison Commelinales 5 families, 780 species, widespread in various habitats Not required 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 (diploid nutritive tissue derived from the nucellus) 8 families and nearly 2000 species Must be able to recognize the order! Zingiberales diversity Musaceae Musa