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The Monocots: Part 2 Commelinid Monocots Spring 2012 Phylogeny of Monocot Groups Basal “Petaloid” Commelinid Acorales Alismatales Asparagales Liliales Dioscoreales Pandanales Arecales Poales Commelinales Zingiberales Fig. 7.17 Commelinid 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 Fig. 7.45 Commelinoid Monocot Groups Order Arecales – Palms Arecaceae (Palmae) Order Commelinales – Spiderworts, bloodworts, pickerel weeds Order Zingiberales – Ginger, banana, and allies Order Poales – Bromeliads, Cat-tails, Rushes, Sedges, and Grasses Typhaceae Juncaceae Cyperaceae Poaceae (Gramineae) Commelinoid Monocots: Arecales: Arecaeae (Palmae) • • • • • • • Widespread throughout tropical and warm temperate regions “Trees” or “shrubs”, typically unbranched Diversity: ca. 2,000 in 190 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 Family not required 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 – Cocos nucifera Arecaceae Economic plants and products: Cocos nucifera Coconut, oil Arecaceae Economic plants and products: Phoenix dactylifera Dates Phylogeny of Monocot Groups Basal “Petaloid” Commelinid Acorales Alismatales Asparagales Liliales Dioscoreales Pandanales Arecales Poales Commelinales Zingiberales Commelinales 5 families, ca. 780 species, widespread in various habitats Not required Commelinid 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 penni-parallel 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 Fig. 7.55 Zingiberales diversity Fig. 7.56 Musaceae Musa Phylogeny of Monocot Groups Basal “Petaloid” Commelinid Acorales Alismatales Asparagales Liliales Dioscoreales Pandanales Arecales Poales Commelinales Zingiberales Characters of Poales • Silica bodies (in silica cells) in the epidermis • Styles strongly branched • Loss of raphide (needle-like) crystals in most • Much molecular support for monophyly • Wind pollination has evolved several times independently within the order • Ecologically very important Fig. 7.63 Commelinid Monocots—Poales: Bromeliaceae (The Pineapple/Bromeliad Family) • Tropical to temperate regions of the Americas • Predominantly epiphytic herbs (“tank” plants) • Diversity: ca. 2,400 species in 59 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 Bromeliaceae: Tillandsia (Spanish moss) Bromeliaceae – Ananas comosus Fruit type? Commelinoid Monocots—Poales: Typhaceae (The Cattail Family) • Widely distributed, especially in Northern Hemisphere • Emergent aquatic rhizomatous herbs • Diversity: 8-13 species in 1 genus • Flowers: small, unisexual; separated • • • spatially on dense, compact spicate inflorescences; placentation apical Significant features: rhizomatous; long slender leaves; characteristic inflorescence Special uses: ornamental aquatics Required taxa: Typha Typha Sparganium This genus is placed in its own family, the Sparganiaceae, in your text, but it is closely related to Typhaceae and is included in Typhaceae in many treatments. Commelinid Monocots—Poales: Juncaceae (The Rush Family) • • • • • • • Worldwide, mostly temperate regions; wet or damp habitats Rhizomatous herbs, stems round and solid Diversity: 350 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 Distichia Juncaceae: Juncus -cymose inflorescences -leaf sheaths open -leaf blades flat, grooved, or cylindrical Commelinid Monocots—Poales: Cyperaceae • • • • • • • (The Sedge Family) Worldwide, usually in damp or semi-aquatic sites Rhizomatous herbs, stems usually triangular in cross section and solid Diversity: 5,000 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.” Fig. 7.66D Fig. 7.65 Flowers: •Arranged in spikelets •Reduced •Wind-pollinated flowers •Subtended by bract •Reduced/absent perianth Sedge 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 -ecologically important, especially in wetlands Cyperaceae: Carex Commelinid Monocots—Poales: Poaceae (Gramineae) • • • • • • • (The Grass Family) Cosmopolitan Primarily herbs, often rhizomatous; “trees” in most bamboos; stems are called culms, hollow or solid Diversity: >10,000 species in ca. 650 genera Flowers: small petals 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: family only 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. Poaceae: caryopsis (grain) Zea mays corn or maize Setaria foxtail 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: Bambusoideae 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! •Triangular, solid, not obviously jointed •3 •Terete, solid, not obviously jointed •Leaf sheath Open, ligule •Closed •Open •Inflor: Spikelets •Spikelets •Cymose •Perianth: Lodicules •None or bristles/scales Achene •6 chaffy tepals •Stem terete, hollow, or solid, jointed •Leaf ranks 2 •Fruit: Caryopsis •3 •Capsule “Graminoids” - Comparison Next time: The “Basal” Eudicots…