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10/14/13 1 Diversity And Classification of Flowering Plants: Monocots Michael G. Simpson Monocotyledons (Monocotyledonae, Monocots, Liliidae) Monophyletic group Comprise 22% of all angiosperms Relationships to other basalangiosperms unclear at present aroids, lilies, gingers, orchids, irises, palms, grasses Apomorphies of the Monocots sieve tube plastids with cuneate (wedge- shaped) proteinaceous inclusions atactostele stem vasculature parallel leaf venation single cotyledon NOTE: a trimerous flower merosity is not a Monocot apomorphy

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Diversity And Classification of Flowering Plants:

Monocots

Michael G. Simpson

Monocotyledons ���(Monocotyledonae, Monocots,

Liliidae) •  Monophyletic group •  Comprise 22% of all angiosperms •  Relationships to other “basal” angiosperms

unclear at present •  aroids, lilies, gingers, orchids, irises, palms,

grasses

Apomorphies of the Monocots

•  sieve tube plastids with cuneate (wedge-shaped) proteinaceous inclusions

•  atactostele stem vasculature •  parallel leaf venation •  single cotyledon

•  NOTE: a trimerous flower merosity is not a Monocot apomorphy

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•  No monocot has true vascular cambium that produces wood

Apomorphies of the Monocots •  sieve tube plastids with cuneate (wedge-shaped)

proteinaceous inclusions

Apomorphies of the Monocots •  atactostele stem vasculature

Apomorphies of the Monocots •  parallel leaf venation (penni-parallel in some)

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Apomorphies of the Monocots •  single cotyledon

ALISMATALES Alismataceae

Aponogetonaceae Araceae (including Lemnaceae) Butomaceae Cymodoceaceae Hydrocharitaceae Juncaginaceae Limnocharitaceae Posidoniaceae Potamogetonaceae Ruppiaceae Scheuchzeriaceae Tofieldiaceae Zosteraceae

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ALISMATALES Araceae - Arum family

(Arum, a name used by Theophrastus). 111 genera / 2980 species

The Araceae are distinguished from related families in having bifacial leaves, a spadix of numerous, small flowers with a subtending spathe, endospermous seeds, and raphide crystals.

P 2+2,3+3,(2+2),(3+3) or 0 [4+4,(4+4)] A 4,6,8 or

(4,6,8) [1-12] G (3) [1-(∞)] superior, hypanthium absent.

Araceae - Arum family •  Members of the family have distributions in

tropical and subtropical regions. •  Economic importance includes many taxa

important food sources (rootstocks, leaves, or fruits) in the tropics, e.g., Alocasia, Colocasia esculenta (taro), Monstera; some species are cultivated ornamentals, e.g., Anthurium, Caladium (elephant's ear), Dieffenbachia (dumb cane), Monstera, Philodendron, Zantedeschia (calla-lily).

•  Amorphophallus titanum is unique in having the largest inflorescences of any flowering plant.

ARACEAE •  Colocasia “gabi” or “taro” •  Alocasia •  Caladium “Corazon de Maria” •  Dieffenbachia “dumb cane” •  Anthurium

Aglaonema commutatum A. modestum

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Amorphophallus titanum Largest inflorescence in world Arisaema triphyllum Jack-in-the-pulpit

Lemna minuscula Duckweed

Wolffia sp. Watermeal Smallest flower in world

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Pistia stratiotes Water-lettuce Monstera deliciosa

Monstera deliciosa

Monstera deliciosa

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Pothos sp. Xanthosoma sagittifolium

Xanthosoma sagittifolium

Zantedeschia aethiopica Calla-lily

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Zantedeschia aethiopica Calla-lily

ASPARAGALES •  Very large order! •  Apomorphy: phytomelan in seed coat���

(lost in some taxa)

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ASPARAGALES Agapanthaceae* [Alliaceae] Agavaceae* [Asparagaceae] Alliaceae* Amaryllidaceae [Alliaceae] Aphyllanthaceae* [Asparagaceae] Asparagaceae Asphodelaceae* [Xanthorrhoeaceae] Asteliaceae Blandfordiaceae Boryaceae Doryanthaceae Hemerocallidaceae* [Xanthorrhoeaceae]

Hyacinthaceae* [Asparagaceae] Hypoxidaceae Iridaceae Ixioliriaceae Lanariaceae Laxmanniaceae* [Asparagaceae] Orchidaceae Ruscaceae* [Asparagaceae] (incl. Convallariaceae) Tecophilaeaceae Themidaceae* [Asparagaceae] Xanthorrhoeaceae* Xeronemataceae

Orchidaceae - Orchid family (orchis, testicle, from the shape of the root tubers). 700-800

genera / ca. 20,000 species •  mycorrhizal, mostly perrennial, terrestrial or epiphytic

herbs having trimerous, •  often resupinate flowers with a showy labellum •  androecium and gynoecium adnate (termed a column,

gynostegium, or gynostemium) •  the pollen grains often fused into a 1-several masses

(pollinia), bearing a sticky-tipped stalk, pollinia and stalk termed a pollinarium, which is the unit of pollen dispersal during pollination.

P (3+3) A 1-3, when 1 a pollinarium G (3), inferior, with gynostemium

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Caladenia longiclavata Australia Calopogon - a non-resupinate orchid

labellum

column

Cattleya sp. Cymbidium sp.

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Cypripedium sp. Lady’s Slipper

Dendrobium phalaenopsis

Dracula chimaera Dracula vampira Encyclia sp. Cockleshell Orchid

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Epidendrum sp. Ludisia sp.

Oncidium lanceanum

Orchis spectabilis

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Paphiopedilum sp.

Pseudobulb

Stanhopea trigrinum

Thelymitra antennifera Australia

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Vanilla planifolia- fermented capsules Vanilla Orchid

Commelinid Monocots Apomorphy: ���

UV-fluorescent organic acids (including coumaric, diferulic, and ferulic acid) that impregnate the cell walls.

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ARECALES ���Arecaceae (Palmae) - Palm family

(from areca, Portuguese for the betel palm). ca. 190 genera / ca. 2,000 species. The Arecaceae are distinctive in having a rhizomatous, lianous, or

usually arborescent stem, with large, sheathing, plicate leaves, a fleshy, usually drupaceous fruit, and seeds lacking starch. The plicate leaves is an apomorphy for the family.

P 3+3 [0,2+2,∞] A 3+3 or (3+3) [3,∞; 0 in female fls.] G 3 or (3) [1,2,4-10; 0 in male fls.], superior.

Arecaceae

•  Leaves- acrocaulis; simple, pinnate, bipinnate, costapalmate, palmate

•  Pseudopetiole •  Crownshaft •  Rhizomatous, lianous, herbs, trees

reduplicate vs. induplicate Fruit a drupe

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•  Cocos nucifera- coconut •  Phoenix dactylifera- date palm •  Elaeis oleifera- oil palm •  Metryoxylon- sago palms •  Copernicia cerifera- wax palm •  Areca catechu- as a stimulant

Archontophoenix cunnighamiana King Palm

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Syagrus romanzoffianum Queen Palm Calamus Rattan Palm

Chamaerops humilis Mediterranean Palm Jubea chilensis

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Licuala peltata Nypa fruticans - a rhizomatous palm

Phoenix canariensis Canary I. Palm

Phoenix dactylifera Date Palm

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Rhopalostylis sapida

ZINGIBERALES - Ginger Group

Apomorphies: Penni-parallel venation Supervolute leaf “ptyxis” (1/2 of leaf rolled within the other)

Diaphraghmed air chambers Silica cells Inferior ovary

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midrib

Zingiberaceae - Ginger family (from a pre-Gr. name, possibly from India). 50 genera / 1,200 species

The Zingiberaceae are distinguished from related families of the Zingiberales in having distichous, usually ligulate leaves with a single, dithecal stamen and a petaloid labellum derived from two staminodes.

P (3+3) A 1 fertile + 2 + (2) petaloid staminodes G (3), inferior.

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Zingiberaceae - Ginger family (from a pre-Gr. name, possibly from India). 50 genera / 1,200 species

Economic importance includes the source of important spice plants, e.g., Curcuma spp., including C. domestica (turmeric), Elettaria cardamomum (cardamom), and Zingiber spp., including Z. officinale (ginger); some species are grown as cultivated ornamentals, e.g., Alpinia and Hedychium.

Zingiberaceae

Strelitziaceae Musaceae

ZINGIBERACEAE •  1 style, clasped

by anther (like Schistosoma)

•  Fruit capsule

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Alpinia Shell Ginger

Outer tepal

inner tepal

Connate staminodes

Hedychium Hedychium

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Hedychium

Ovary

staminodes

Outer tepals

Inner tepals

filament

Nicolaia elatior

Cannaceae - Canna-Lily family (Gr. canna, a reed). 1 genus (Canna) / 25 species

The Cannaceae are distinguished from related families of the Zingiberales in having usually distichous leaves and flowers with one petaloid, monothecal stamen associated with 1-4[5] petaloid staminodes.

P 3+3 A 1, petaloid & monothecal + 1-4 petaloid

staminodes G (3), inferior.

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Cannaceae

Musaceae

Canna sp. Cannaceae

Canna sp. Cannaceae

Outer tepal

Inner tepal

staminodes

Canna sp. Cannaceae

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Canna sp. Cannaceae

staminode half staminode

style/stigma

half anther

•  Fruit capsule •  Canna sp.

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Cyperaceae - Sedge family (Gr. for several species of Cyperus). 98 genera / 4,350 species

The Cyperaceae are distinctive in being herbs with usually 3-sided, solid-pithed stems, closed-sheathed, often tristichous leaves, the inflorescence a “sedge spikelet,” consisting of a central axis bearing many sessile, distichous or spiral bracts, each subtending a single, reduced unisexual or bisexual flower, with perianth absent or reduced to bristles or scales, usually 3 stamens, and a 2–3-carpellate ovary, the fruit a 2- or 3-sided achene. P 6 or 0 [1-∞] A 3 [1-6+] G (2-3)[(4)], superior.

Cyperaceae - Sedge family (Gr. for several species of Cyperus). 98 genera / 4,350 species

Economic importance is limited, with some species used as mats, thatch, weaving material, or writing material (Cyperus papyrus, papyrus, the culm pith of which was historically used to make paperlike scrolls), a few used as ornamental cultivars (e.g., Cyperus involucratus, umbrella plant), and some species, such as the nutsedges, being noxious weeds.

Glomerule

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Cyperus involucratus Umbrella Sedge

spikelet involucral

bracts

Carex praegracilis

female spikelet

perigynia enclosing ovaries

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Cyperus sp.

spikelet: bracts

distichous

Cyperus involucratus

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Poaceae/ Graminae - Grass family (after Poa, Gr. name for a grass). 668 genera / 9,500 species

STEMS: hollow-pithed LEAVES: open-sheathed, distichous, with a ligule at inner

junction with blade INFLORESCENCE: grass spikelet

= axis + two basal glumes + 1-∞ florets Floret = short lateral axis + lemma + palea + flower

FLOWER: P 2-3 lodicules A 2-3 (pendulous) G (2-3) FRUIT: caryopsis (grain).

POACEAE

•  Spikelet – basic inflorescence, made up of florets, subtended by 2 bracts called glumes

•  Floret – flower, each subtended by even more glumes called lemma and palea, uni- or bisexual

•  Rachilla – axis of spikelet

POACEAE

•  Lemma – outer, flowering glume •  Palea – inner bract, between flower and

rachilla •  Lodicules – modified perianth •  Stamens in 3 whorls

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DISTRIBUTION: worldwide. ECONOMIC IMPORTANCE:

1) agricultural grains (most important food in world): barley (Hordeum) corn (Zea) oats (Avena) rice (Oryza) rye (Secale) wheat (Triticum); 2) alcoholic beverages: beer, whiskey, gin 3) forage and grazing plants; 4) important components of many ecosystems, such as grasslands and savannahs.

POACEAE •  Oryza sativa “rice” •  Zea mays “corn” •  Triticum “wheat” •  Avena “oats” •  Hordeum “barley” •  Saccharum “sugar cane” •  Panicum •  Axonopus •  Eleusine •  Digitaria •  Paspalum

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distichous leaf

arrangement

open sheath

ligule

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Bamboos Fruit: Grain

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Cynodon dactylon Bermuda Grass

Pennisetum setaceum

Melinis repens