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Diversity And Classification of Flowering Plants:

Eudicots

Michael G. Simpson

Eudicots

•  Palynological apomorphy: tricolpate or tricolpate-derived pollen grain

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Eudicots UNPLACED** Apodanthaceae, Balanophoraceae, Cynomoriaceae, Cytinaceae, Hoplestigmataceae, Medusandraceae, Metteniusaceae, Pottingeriaceae BUXALES**

Buxaceae Didymelaceae**

Sabiaceae TROCHODENDRALES**

Trochodendraceae RANUNCULALES

Berberidaceae Circaeasteraceae Eupteleaceae Lardizabalaceae Menispermaceae Papaveraceae Ranunculaceae

PROTEALES Nelumbonaceae Platanaceae* [Proteaceae] Proteaceae

CORE EUDICOTS BERBERIDOPSIDALES**

Aextoxicaceae Berberidopsidaceae

DILLINEALES** Dilleniaceae

GUNNERALES Gunneraceae Myrothamnaceae**

CARYOPHYLLALES Achatocarpaceae Aizoaceae Amaranthaceae Ancistrocladaceae Asteropeiaceae Barbeuiaceae Basellaceae Cactaceae Caryophyllaceae Didiereaceae Dioncophyllaceae

Droseraceae Drosophyllaceae Frankeniaceae Gisekiaceae Halophytaceae** Hectorellaceae** Limeaceae** Molluginaceae Nepenthaceae Nyctaginaceae Physenaceae Phytolaccaceae Plumbaginaceae Polygonaceae Portulacaceae Rhabdodendraceae Sarcobataceae Simmondsiaceae Stegnospermataceae Tamaricaceae

Eudicots (continued) SANTALALES

Erythropalaceae** Olacaceae Opiliaceae Loranthaceae Misodendraceae Santalaceae Schoepfiaceae**

SAXIFRAGALES Altingiaceae Aphanopetalaceae** Cercidiphyllaceae Crassulaceae Daphniphyllaceae Grossulariaceae Haloragaceae Hamamelidaceae Iteaceae Paeoniaceae Penthoraceae** Peridiscaceae** Pterostemonaceae** Saxifragaceae Tetracarpaeaceae**

ROSIDS (see Table 8.2) ASTERIDS (see Table 8.3)

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Diversity And Classification of Flowering Plants:

Eudicots: Rosids

Michael G. Simpson

ROSIDS

•  Very large, monophyletic group of Eudicots •  Linked by no clear non-molecular

apomorphies •  Ovules bitegmic (2 integuments) &

crassinucellate [contrast with Asterids] •  13 orders

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Aphloiaceae Geissolomataceae Ixerbaceae Picramniaceae Strasburgeriaceae Vitaceae CROSSOSOMATALES Crossosomataceae Stachyuraceae Staphyleaceae GERANIALES Geraniaceae (incl. Hypseocharitaceae) Ledocarpaceae Melianthaceae (incl. Francoaceae) Vivianiaceae

MYRTALES Alzateaceae Combretaceae Cryteroniaceae Heteropyxidaceae Lythraceae Melastomataceae (incl. Memecylaceae) Myrtaceae Oliniaceae Onagraceae Penaeaceae Psiloxylaceae Rhynchocalycaceae Vochysiaceae

EUROSIDS I Zygophyllaceae Krameriaceae* Huaceae CELASTRALES Celastraceae (incl. Lepidobotryaceae) Huaceae Parnassiaceae (incl. Lepuropetalaceae) CUCURBITALES Anisophylleaceae Begoniaceae Coriariaceae Corynocarpaceae Cucurbitaceae Datiscaceae Tetramelaceae

EUROSIDS I (continued) FABALES Fabaceae (Leguminosae) Polygalaceae Quillajaceae Surianaceae FAGALES Betulaceae Casuarinaceae Fagaceae Juglandaceae (incl. Rhoipteleaceae) Myricaceae Nothofagaceae Ticodendraceae

MALPIGHIALES Achariaceae Balanopaceae Bonnetiaceae Caryocaraceae Chrysobalanaceae (incl. Dichapetalaceae, Euphroniaceae,

Trigoniaceae) Clusiaceae (Guttiferae) Ctenolophonaceae Elatinaceae Erythroxylaceae* Euphorbiaceae, s.l. Goupiaceae Humiriaceae Irvingiaceae Ixonanthaceae Lacistemataceae

MALPIGHIALES (continued) Linaceae Lophopyxidaceae Malpighiaceae Ochnaceae (incl. Medusagynaceae, Quiinaceae) Pandaceae Passifloraceae (incl. Malesherbiaceae, Turneraceae) Peridiscaceae Phyllanthaceae Picrodendraceae Podostemaceae Putranjivaceae Rhizophoraceae Salicaceae Violaceae

EUROSIDS I (continued) OXALIDALES Brunelliaceae Cephalotaceae Connaraceae Cunoniaceae Elaeocarpaceae Oxalidaceae ROSALES Barbeyaceae Cannabaceae Dirachmaceae Elaeagnaceae Moraceae Rhamnaceae Rosaceae Ulmaceae Urticaceae

EUROSIDS II Tapisciaceae BRASSICALES Akaniaceae [incl.

Bretschneideraceae] Bataceae Brassicaceae (Cruciferae) Caricaceae Emblingiaceae Gryrostemonaceae Koeberliniaceae Limnanthaceae Moringaceae Pentadiplandraceae Resedaceae Salvadoraceae Setchellanthaceae Tovariaceae Tropaeolaceae

MALVALES Bixaceae Cistaceae Dipterocarpaceae Malvaceae Mutingiaceae Neuradaceae Sarcolaenaceae Sphaerosepalaceae Thymelaeaceae SAPINDALES Anacardiaceae Biebersteiniaceae Burseraceae Kirkiaceae Meliaceae Nitrariaceae Rutaceae Sapindaceae Simaroubaceae

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Myrtaceae - Myrtle family (myrtus, Gr. name for myrtle). 120 genera / 3850 species

•  trees and shrubs •  with glandular-punctate or pellucid leaves •  Usually epiperigynous flowers with numerous

stamens. K 4-5 [3,6] C 4-5 [3,6] A ∞ G (2-5) [(-16)],

inferior [rarelyhalf-inferior or superior], with hypanthium.

Myrtaceae - Myrtle family (myrtus, Gr. name for myrtle). 120 genera / 3850 species

Timber, oil - Eucalyptus spp. edible fruits - Psidium guajava, guava spices - Syzygium aromaticum, cloves, cultivated ornamentals -Callistemon (bottlebrush),

Eucalyptus sideroxylon

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Leptospermum laevigatum

FABALES Fabaceae (Leguminosae)

- Bean/Pea family (after faba, Latin name for broad bean). 643 genera / 18,000 species •  trees, shrubs, vines, or herbs, •  stipulate, often compound leaves •  Single, unicarpellous pistil with marginal

placentation •  legume (or modified legume) K 5 or (5) [(3-6)] C 5 or (5) [0,1-6, or (1-6)] A

10 or (10) to ∞ [variable] G 1 [2-16], superior, hypanthium sometimes present

worldwide distribution ecologically important for nitrogen-fixing

rhizobial nodules pulses (e.g., beans, peas, soybeans, etc.),

fodder plants, oils, timber trees, gums, dyes, and insecticides.

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Fabaceae: 3 subfamilies Caesalpinioideae Flowers zygomorphic; petals distinct; posterior petal inner

to laterals; stamens distinct. Mimosoideae Flowers actinomorphic; petals distinct or connate; stamens

often ∞, showy; flowers often densely aggregated. Faboideae (=Papilionoideae) Flowers zygomorphic; perianth papilionaceous; posterior

petal outer to laterals; stamens connate.

Caesalpinioideae

Flowers zygomorphic Petals distinct Posterior petal inner to laterals Stamens distinct

Bauhinia variegata Orchid Tree

posterior petal inner to laterals

Bauhinia variegata Orchid Tree

stamens distinct

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Bauhinia variegata Orchid Tree

ovary

style

stipe

Cassia alata - akapulko

Purgative, for cough, fungicide���stomatitis.

���

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CAESALPINOIDEAE

•  Caesalpinia pulcherrima – “caballero” •  Caesalpinia inerma •  Cassia fistula – “golden shower” •  Cassia alata – “Acapulco”

CAESALPINOIDEAE

•  Delonix regia – “flame tree” •  Bauhinia purpurea – “alibangbang” •  Cynometra •  Tamarindus indica •  Intsia – “ipil”

Flowers actinomorphic, often densely aggregated Petals distinct or connate; hypanthium sometimes

present Stamens often ∞, showy

Mimosoideae

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Acacia spp.

heads

Acacia spp.: phyllodinous

phyllode

rachillae ���with leaflets

phyllode

Acacia longifolia (native to Australia)

spike

ovary (removed)

flowers actinomorphic, stamens ∞

Calliandra haematocephala

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Mimosa sp. Pithecellobium unquis-cati Cat Claw

MIMOSOIDEAE

•  Mimosa pudica – “makahiya” •  Samanea saman – “acacia” •  Acacia sp. •  Leucaena leucocephala – “ipil-ipil” •  Calliandra

MIMOSOIDEAE

•  Enterolobium cyclocarpum •  Pithecellobium dulce – “camachile” •  Albizia •  Adenanthera •  Parkia roxburghii

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Flowers zygomorphic Perianth papilionaceous Terminology: Posterior petal = banner or standard Lateral petals = wings Anterior petals = keel petals (basally distinct; distally connate; collectively called the keel) Posterior petal (banner) outer to laterals (wings)

Stamens connate: monadelphous or diadelphous

Faboideae (Papilionoideae)

PAPILIONOIDEAE

•  Phaseolus – “abitsuelas”, “lima bean”, “patani”

•  Vigna – “munggo”, “sitaw” •  Psophocarpus – “sigarillas” •  Sesbania – “katuray”

PAPILIONOIDEAE

•  Dolichos lablab – “batao” = Lablab purpureus

•  Pterocarpus indica – “narra” •  Arachis pintoi – “ornamental peanut” •  Arachis hypogea – “peanut”

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PAPILIONOIDEAE

•  Erythrina subumbrans – “dapdap” •  Pisum sativum – “sitsaro”, “peas” •  Centrosema •  Clitorea •  Desmodium

PAPILIONOIDEAE

•  Gliricidia – “kakawate” or “madre de cacao”

•  Crotalaria (with inflated pods) •  Mucuna pruriens – “lipa” •  Pachyrrizus erosus – “singkamas” •  Abrus precatorius

Wisteria sinensis Wisteria

banner outer to laterals

wing petals

keel

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Wisteria sinensis Wisteria

keel petal

stamens connate: diadelphous (9+1)

in Wisteria

Wisteria sinensis Wisteria

style (ovary hidden)

Erythrina caffra

banner

wing petals

stamens calyx

Erythrina caffra

stamens connate: diadelphous (9+1)

style

pistil removed stipe

ovary style

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Erythrina caffra

pistil unicarpellous placentation

marginal

(l.s.)

(c.s.)

Clitoria mariana -a resupinate papilionoid

Euphorbiaceae - Spurge family (after Euphorbus, physician to the king of Mauritania, 1st century).

313 genera / 8,100 species unisexual flowers with a superior, usually 3-carpellate ovary with 1 ovule per carpel, apical-axile in placentation;

Crotonoideae and Euphorbioideae have a red, yellow, or usually white (“milky”) latex Euphorbioideae -cyathium inflorescence.

K 5 [0] C 5 [0] A 1-∞ G (3) [(2–∞)], superior.

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Three subfamilies:������Acalyphoideae������Crotonoideae��� -colored latex������Euphorbioideae��� - milky (white) latex��� - inflorescence a cyathium

cyathium

An inflorescence bearing small, unisexual flowers and subtended by an involucre (frequently with petaloid glands), the entire inflorescence resembling a single flower.

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EUPHORBIACEAE

•  Manihot esculenta “cassava” or “kamoteng kahoy”

•  Euphorbia spp. •  Euphorbia pulcherrima “poinsettia” •  Jatropha spp. •  Ricinus communis “castor plant” •  Antidesma “bignay”

EUPHORBIACEAE

•  Phyllanthus acidus “karmay” •  Acalypha hispida “chenille plant” •  Pedilanthus tithymalloides •  Aleurites moluccana “lumbang”,

“candlenut tree”

•  Ricinus communis, the source of castor bean oil and the deadly poison ricin

•  Hevea brasiliensis, the major source of natural rubber;

•  Manihot esculentus, cassava/manioc

•  Oil (biodiesel, Jatropha), timber, medicinal, dye, and ornamental plants.

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Euphorbia grandicornis

Euphorbia millii

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Euphorbia spp.

Manihot esculenta Manioc

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Moraceae — Mulberry family ���(Latin name for mulberry). ca. 40 genera / 1100 species ���

monoecious or dioecious trees, shrubs, lianas, or

herbs milky latex, stipulate, simple leaves fruit a multiple of achenes or syconium. P (0-10) A 1-6 G (2) [(3)], superior or inferior.

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MORACEAE

•  Ficus sp. – figs •  Artocarpus – “jackfruit”, “marang”,

“antipolo” •  Broussonetia “himbabao” •  Morus alba “mulberry”

Artocarpus altilis (breadfruit), Ficus carica (edible fig), Morus spp. (mulberry);

paper, rubber, and timber trees; and some cultivated ornamentals, -Ficus spp., figs;

the leaves of Morus alba are the food source of silkworm moth larvae.

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Rosaceae - Rose family (Latin for various roses). 95 genera / 2,800 species

The Rosaceae are distinctive in having usually stipulate leaves (often adnate to petiole) and an actinomorphic, generally pentamerous flower with hypathium present, variable in gynoecial fusion, ovary position, and fruit type.

K 5[3-10] C 5[0,3-10] A 20-∞[1,5] G 1-∞,

superior or inferior, hypanthium present.

Rosoideae Apocarpous Achenetum/ drupecetum

Fragaria, Rosa

Prunoideae Single pistil; superior

drupe Prunus “peach”

Maloideae Inferior ovary

pome apple

ROSOIDEAE

•  Carpels numerous, fruit achene or drupelet, cyanogenic glycosides absent

•  Rubus “raspberry” “blackberry” •  Rosa •  Fragaria vesca

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PRUNOIDEAE

•  Carpel usually 1, fruit a drupe, with HCN •  Prunus “peach”, “plum”, “prunes”,

“apricot”, “cherry”, “almonds”

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MALOIDEAE

•  Carpels usually 2-5, connate or adnate to hypanthim, fruit a pome, with HCN

•  Malus “apple” •  Pyrus communis “pear”

Rosaceae economically important

Fragaria (strawberry)

Malus (apples)

Prunus (almond, apricot, cherry, peach, plum)

Pyrus (pear)

Rubus (blackberry, raspberry)

ornamentals

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Fragaria vesca Strawberry

Malus pumila Apple

hypanthium inferior ovary

Prunus spp. Cherries, Peaches, Plums

Rosa spp.

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Brassicales

Glucosinolates -. - deter herbivory and parasitism - flavoring agents in brocolli, cauliflower, and

mustard.

Brassicaceae (Cruciferae) — Mustard family ���(name used by Pliny for cabbagelike plants). ���

365 genera / 3250 species. •  with glucosinolates (mustard oil glucosides), •  the perianth cruciate (petals usually clawed) •  androecium with usually 2+4, tetradynamous stamens •  the gynoecium with a superior, 2- carpellate/loculate

ovary •  with axile-parietal placentation •  dehiscent fruit with a replum (silique or silicle). K 2+2 C 4 A 2+4 [2,4-16] G (2), superior.

Brassica oleracea - broccoli, brussels sprouts, cauliflower, cabbage,

collards, kale

rutabaga and canola oil (B. napus),

mustard (B. nigra),

turnip (B. rapa),

Arabidopsis thalliana

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BRASSICACEAE

•  Brassica oleracea “cabbage”, “cauliflower”, “broccoli”, “kohlrabi”, “turnip”, “Brussels sprouts”

•  Brassica juncea “mustasa” •  Raphanus sativus “radish” •  Nasturtium “watercress” •  Rorippa •  Arabidopsis thaliana

Arabidopsis thaliana ���The Model Plant

Brassica nigra Raphanus sativus

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Malvaceae, s.s. - Mallow family (name used by Pliny, meaning "soft"). 111 genera / 1,800 species

Malvaceae s. l. •  herbs, shrubs, or trees, •  often with stellate trichomes, •  typically with an epicalyx, •  calyx valvate, corolla often convolute •  stamens connate into tube or 5-∞ bundles •  Monothecal or bithecal anthers •  K 3-5 or (3-5) C 5 [3- or 0] A 5-∞ G 2-∞ [1],

superior [rarely inferior].

A.P.G.: Malvaceae, s.l.���formerly 4 families:

Malvaceae, s.s. Bombacaceae Sterculiaceae Tiliaceae

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Economic importance includes medicinal plants; Gossypium spp. (cotton, the world’s most important fiber plant) Ceiba pentandra (kapok), in both of which the seed trichomes are

utilized, Corchorus spp. (jute), a bast fiber plant food and flavoring plants -  Theobroma cacao (cacao, the source of chocolate), Cola nitida

(cola), Abelmoschus (okra), and Durio zibethinus (durian); -  ornamental cultivars such as Chorisia (floss-silk tree), Hibiscus

(mallows), and Tilia (linden tree). -  Adansonia digitata (baobab, tropical Africa) - economic or

ecological importance.

Apomorphies of Malvaceae Inflorescence with “bicolor unit” (after

Theobroma bicolor), consisting of a modified, 3-bracted cyme, the trimerous epicalyx of family members possibly derived from these 3 bracts.

Other apomorphies: valvate calyx stellate or lepidote trichomes, dilated secondary tissue rays

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Hibiscus sp. Kosteletskia virginica

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Bombax glabrum

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Chorisia speciosa Chorisia speciosa

Chorisia speciosa Durio

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Theobroma cacao Cacao, source of

chocolate