The Monocots: Part 1 Overview, Basal, and “Petaloid” Groups Spring 2012 Figure 7.1 from the text Synapomorphies of Monocots • Root system adventitious • One cotyledon • Stems with scattered vascular bundles (no • • • • secondary growth); herbaceous Leaves parallel-veined with a sheathing base Flowers pentacyclic (5 whorls), trimerous Sieve cell plastids with several cuneate protein crystals Lots of molecular support for monophyly Additional features of monocots • Leaves formed from the basal end of the leaf primordium • Usually with monosulcate pollen • Lack glandular teeth on leaves Monocot characters • One cotyledon! NONMONOCOT MONOCOT Monocot characters Leaves: – parallel venation in most monocots [may be reversals with net-venation!] – sheathing base Trillium Smilax Monocot characters Cuneate protein bodies in sieve cell plastids – “wedge-shaped” inclusions – function unknown Monocot characters Adventitious roots: -derived from structures other than another root Monocot characters Scattered vascular bundles in stem – numerous; actually complex organization – no vascular cambium (a few weird exceptions) Monocot characters • Pentacyclic, trimerous flowers with 2 perianth whorls and two whorls of stamens How many monocots? •ca. 3,000 genera •ca. 65,000 species •22-25% of angiosperms •Include: -aroids -bananas -lilies -gingers -orchids (20,000+ spp.) -irises -palms -grasses (10,000 spp.) Fig. 7.17 from Simpson Phylogeny of Monocot Groups Basal “Petaloid” Commelinoid Acorales Alismatales Liliales Asparagales Dioscoreales Pandanales Arecales Poales Commelinales Zingiberales Basal and “Petaloid” Monocot Groups Order Acorales Acoraceae Order Alismatales Araceae Alismataceae Order Liliales Liliaceae Order Asparagales Agavaceae Alliaceae Amaryllidacaee Iridaceae Orchidaceae Basal Monocots: Acorales: Acoraceae • Widespread, temperate throughout tropical regions • Aquatic herb • Diversity: 1-3 spp. in 1 genus (Acorus) • Flowers: typical of Araceae, coalesced into a spike-like spadix • Significant features: Sister to the rest of the monocots; contain ethereal oils. • Special uses: none • Family not required, but Acorus evolutionarily important Acorus (sweet flag)– The most basal monocot! Aquatic. “Petaloid” Monocots—Alismatales: Araceae (The Arum Family) • • • • • • • Cosmopolitan; greatest diversity in tropical regions Terrestrial and aquatic herbs, vines, epiphytes, floating aquatics Diversity: 3,300 species, 109 genera Flowers: many, small; lacking extensive perianth, carpels 2-3; if unisexual then spatially separated in inflorescence or sometimes plants dioecious Significant features: inflorescence – spadix subtended by a spathe (specialized leaf) Special uses: many ornamentals; Colocasia as food Required taxa: Arisaema, Lemna Araceae—Arisaema Arisaema triphyllum Jack-in-the-pulpit Arisaema dracontium green dragon Arisaema sikokianum -spathe margins overlapping below, spathe mostly arched above, striped or marked -spadix usually slender and elongate -flowers unisexual and only at the base of the spadix Araceae Philodendron Monstera Amorphophallus Araceae: Lemna and friends •Reduced plant body: no stem or leaves; sometimes no roots •Rarely flower Lemna ~ duckweed Alismatales: Araceae Economic plants and products: Colocasia esculenta •Taro “root” or dasheen “poi” •10% of the world uses as staple (starch) in diet “Petaloid” Monocots—Alismatales: Alismataceae (The Water Plantain Family) • Widely distributed • Aquatic & wetland rhizomatous herbs • Number of species: 88 species, 15 genera • Flowers: sepals & petals distinct, many • • • apocarpous carpels; flowers or floral axes often whorled Significant features: rhizomatous Special uses: ornamental aquatics Family not required Phylogeny of Monocot Groups Basal “Petaloid” Commelinoid Acorales Alismatales Liliales Asparagales Dioscoreales Pandanales Arecales Poales Commelinales Zingiberales Liliales • Nectaries at base of tepals • Spots on tepals • Extrorse anthers “Petaloid” Monocots—Liliales: Liliaceae (The Lily Family) • Widely distributed in temperate regions of the Northern Hemisphere • Perennial herbs, usually with bulbs and contractile roots • Number of species: ca. 600 species, in 16 genera • Flowers: tepals 6, distinct, carpels 3, stamens 6 • Significant features: Fruit a loculicidal • • capsule, sometimes a berry; no onion-like odor Special uses: many ornamentals Required taxa: Erythronium, Tulipa Liliaceae - Lilium Liliaceae Erythronium trout-lily -bulbs ovate to elongate -scapose herbs with 2 leaves (1 if non-flowering) -tepals 6, spreading to reflexed -native wildflowers Tulipa -scapose herbs from tunicate bulbs -leaves 2-several on a stem -perianth campanulate to cuplike -tepals 6, erect -stigma prominently 3-lobed Liliaceae Economic plants and products (horticultural): Lilium Easter lily Tulipa tulip Phylogeny of Monocot Groups Basal “Petaloid” Commelinoid Acorales Alismatales Liliales Asparagales Dioscoreales Pandanales Arecales Poales Commelinales Zingiberales Asparagales vs. Liliales • Herbs to woody; • Herbs; not succulent sometimes succulent •Tepals not spotted •Nectaries septal •Style usually 1, simple •Seed coat collapsed to + present •Phytomelan crust (seeds black) from dry fruits; not in fleshy fruit •Tepals often spotted •Nectaries at base of tepals/filaments •Styles 1 (trifid) or 3 •Seed coat present •No phytomelan crust (seeds not black) Figure 7.32 from the text “Petaloid” Monocots—Asparagales: Alliaceae • • • • • • • (Onion Family) Widely distributed in temperate and tropical regions; also semiarid. Bulb-forming herbs with basal, usually narrow leaves Number of species: ca. 600 species, in 13 genera Flowers: Often showy, tepals 6, stamens 6, 3 connate carpels, ovary superior; inflorescence umbellate; fruit a loculicidal capsule. Significant features: sulfur-containing compounds (onion odor) Special uses: onion, garlic, leek, shallots, chives, used as food & seasonings; ornamentals Required taxa: Allium Alliaceae - Allium -scapose herbs with bulbs + contractile roots -basally clustered leaves -umbellate inflorescence with bracts -6 petaloid tepals + 6 stamens -loculicidal capsule + black seeds Alliaceae Economic plants and products: •Allium species – onions, leeks, garlic! Ornamentals “Petaloid” Monocots—Asparagales: Agavaceae (The Agave Family) • Warm temperate to tropical regions of the New World; maximum diversity in Mexico • Rosette herbs to small trees, often with succulent leaves • Number of species: ca. 300 species in 8-13 genera • Flowers: tepals 6, stamens 6, carpels 3, fruits a loculicidal capsule • Significant features: large, paniculate inflorescence; dimorphic chromosomes • Special uses: fiber, tequila, ornamentals. • Family not required Agavaceae – Agave and Yucca Agave Yucca Agave: bat pollinated Yucca: moth pollinated Agave (L) vs. Yucca (R) • Perianth tubularfunnelform, 6-parted • Stamens exserted beyond the perianth, anthers versatile • Ovary inferior • Capsule loculicidal • Bat-pollinated • Perianth of 6 flat, free tepals • Stamens shorter than the tepals, anthers basifixed • Ovary superior • Fruit indehiscent (berry-like) or septicidal capsule • Moth-pollinated Agavaceae: Hosta -rhizomatous, scapose perennials -leaves with a distinct petiole -perianth tubular-funnelform, white, bluish or lavender -stamens 6, epipetalous or hypogynous -fruit a loculicidal capsule Asparagales: Agavaceae Economic plants and products: Agave tequila Asparagales: Agavaceae Economic plants and products: •Fiber for rope from species of Yucca and Agave e.g., sisal hemp “Petaloid” Monocots—Asparagales: Amaryllidaceae (Amaryllis or Daffodil Family) • • • • • • • Widely distributed in temperate to tropical regions; maximal diversity in South Africa, Andean South America, and the Mediterranean Bulb-forming herbs with contractile roots Number of species: 850 species in 59 genera Flowers: often showy; tepals 6; stamens 6, sometimes adnate to perianth; carpels 3, inferior ovary; fruit usually a loculicidal capsule Significant features: special alkaloid compounds present Special uses: many ornamentals (Narcissus, Hippeastrum) Family not required Amaryllidaceae diversity Amaryllidaceae Corona sometimes present Hymenocallis spider-lily Narcissus daffodil, jonquil, narcissus Narcissus -scapose, perennial herbs from bulbs -perianth of 6 basally connate tepals, yellow and/or white -cuplike to trumpetlike corona present -stamens 6, epipetalous Amaryllidaceae: Hippeastrum -perennial, scapose herbs from large bulbs -perianth of 6 basally connate tepals, white to pink to salmon or red -corona minute -stamens 6, epipetalous “Petaloid” Monocots—Asparagales: Iridaceae • • • • • • • (The Iris Family) Widespread in tropical and subtropical regions; absent in Australia. Perennial herbs forming rhizomes, corms, or bulbs Number of species: ca. 1,750 species, 67 genera Flowers: radial or bilateral, showy; tepals 6, outer tepals often differentiated from inner; stamens (2) 3, opposite outer tepals; carpels 3, fused into an inferior ovary; fruit a loculicidal capsule Significant features: leaves unifacial or terete, equitant Special uses: many ornamentals; saffron (Crocus sativus) Required taxa: Iris Iridaceae diversity Iris -rhizomatous herbs -leaves equitant, in a fan -spathes 2 -style branches broad, petaloid, terminating in paired crests anthers appressed to style branches “Petaloid” Monocots—Asparagales: Orchidaceae (The Orchid Family) • • • • • • • Widespread throughout the world; maximal diversity in tropical regions Primarily epiphytes; some terrestrial herbs, occasionally vines Diversity: ca. 20,000 species in 700-800 genera Flowers: showy, usually resupinate, bilateral, the median inner tepal differentiated into a labellum (lip); highly modified androecial and gynoecial parts, fused into a column; pollen grouped into soft or hard masses (pollinia) united by a stalk into a pollinarium; ovary inferior; placentation parietal; fruit a capsule dehiscing with (1-)3 or 6 slits; seeds tiny, dust-like Significant features: among the most specialized of all angiosperm flowers Special uses: many ornamentals; Vanilla Required taxa: family only Orchid flower morphology pollinarium Pollination •function of column & pollinia Orchidaceae http://www.youtube.com/watch?v=wmgKABRCZpo&feature=related Richard Dawkins talking about orchid pollination Morgan’s Sphinx Moth Endemic to Madagascar Comet Orchid (Angraecum sesquipedale) Asparagales: Orchidaceae Economic plants and products: Vanilla flavoring extracted from immature capsules of Vanilla planifolia