The Monocots: Part 1 Overview, Basal, and “Petaloid” Groups Spring 2014

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 tube member plastids with several cuneate protein crystals Lots of molecular support for monophyly

Monocot characters Adventitious roots: -derived from structures other than another root

Monocot characters One cotyledon! MONOCOT NON- MONOCOT

Monocot characters Scattered vascular bundles in stem numerous; actually complex organization no vascular cambium (a few weird exceptions)

Monocot characters Leaves: parallel venation in most monocots [may be reversals with net-venation!] sheathing base Trillium Smilax

Monocot characters Pentacyclic, trimerous flowers with 2 perianth whorls and 2 whorls of stamens and the gynoecium as 1 whorl

Monocot characters Cuneate protein bodies in sieve cell plastids “wedge-shaped” inclusions function unknown

Additional features of monocots Leaves formed from the basal end of the leaf primordium Usually with monosulcate pollen Lack glandular teeth on leaves

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 (11,000+ spp.)

Fig. 7.17 from Simpson

Phylogeny of Monocot Groups Acorales Alismatales Liliales Asparagales Dioscoreales Pandanales Arecales Poales Commelinales Zingiberales Basal “Petaloid” Commelinid

Basal and “Petaloid” Monocot Groups Order Acorales Acoraceae Order Alismatales Araceae* Alismataceae Order Liliales Liliaceae* Order Asparagales Agavaceae Alliaceae* Amaryllidaceae Iridaceae* Orchidaceae* *required families

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

The most basal monocot! Aquatic. Acorus (sweet flag)– The most basal monocot! Aquatic.

“Petaloid” Monocots—Alismatales: Araceae (The Arum Family) Cosmopolitan; greatest diversity in tropical regions Mainly terrestrial and some aquatic herbs, vines, epiphytes, floating aquatics Diversity: 3,300 species, 104 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 family

Araceae—Arisaema Arisaema dracontium green dragon Arisaema triphyllum Jack-in-the-pulpit Arisaema sikokianum -Jack-in-the-pulpit is one of our common spring wildflowers

Araceae Philodendron Monstera

Amorphophallus (Corpse flower) http://www.youtube.com/watch?v=FHaWu2rcP94

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 Acorales Alismatales Liliales Asparagales Dioscoreales Pandanales Arecales Poales Commelinales Zingiberales Basal “Petaloid” Commelinid

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 family

Liliaceae - Lilium

Liliaceae Erythronium trout-lily -native spring wildflower of woodlands

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): Tulipa Lilium Easter lily

Phylogeny of Monocot Groups Acorales Alismatales Liliales Asparagales Dioscoreales Pandanales Arecales Poales Commelinales Zingiberales Basal “Petaloid” Commelinid

Asparagales vs. Liliales Herbs to woody; 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 Herbs; not succulent 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 family

Alliaceae - Allium

Economic plants and products: Alliaceae Economic plants and products: Allium species – onions, leeks, garlic! Ornamentals

“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 family

Iridaceae characters Perennating structures Equitant leaves Stamen position opposite outer tepals Iridaceae characters

Iridaceae diversity

Iris (Greek for rainbow) -style branches broad, petaloid, terminating in paired crests -anthers appressed to style branches

Iris in wetland habitats

“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 family

Orchid growth habits Epiphytic Terrestrial

Orchid roots velamen

Orchid flower morphology see Digital Flowers

Orchidaceae pollinarium Pollination function of column & pollinia

Richard Dawkins talking about orchid pollination http://www.youtube.com/watch?v=wmgKABRCZpo&feature=related Richard Dawkins talking about orchid pollination

Comet Orchid (Angraecum sesquipedale)

http://www.youtube.com/watch?v=OMVN1EWxfAU Morgan’s Sphinx Moth Endemic to Madagascar

Asparagales: Orchidaceae Economic plants and products: Vanilla flavoring extracted from immature capsules of Vanilla planifolia

Some other cool Asparagales Agavaceae Amaryllidaceae

Agavaceae – Agave and Yucca Hosta Yucca

Agave: bat pollinated Yucca: moth pollinated

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

Amaryllidaceae diversity

Amaryllidaceae Corona sometimes present Hymenocallis Narcissus spider-lily Narcissus daffodil, jonquil, narcissus