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Core

• two well supported lineages of the ‘true’ or core Diversity and Evolution asterids

• ‘lamiid’ or Asterid I group lamiids of Asterids • ‘campanulid’ or Asterid II group

. . . gentians, milkweeds, and • appear to have the typical fused corolla derived independently and via two different floral potatoes . . . developmental pathways campanulids

lamiid campanulid

Core Asterids Early vs. Late Sympetaly euasterids II - campanulids euasterids I - lamiids • two well supported lineages of the ‘true’ or core asterids

lamiids = NOT fused corolla tube

• Asterids primitively NOT fused corolla at maturity campanulids • 2 separate origins of fused in “core” Asterids (plus several times in Ericales) Calendula, Asteraceae early also in Cornaceae of Anchusa, Boraginaceae late ”basal asterids”

1 Gentianales

• order within ‘lamiid’ or Asterid I group

• 5 families and nearly 17,000 dominated by () and (milkweed) lamiids

• iridoids, opposite , contorted corolla

Rubiaceae Apocynaceae campanulids

corolla aestivation

* - gentians *Gentianaceae - gentians Cosmopolitan of 87 genera and nearly 1700 species. Herbs to small • opposite leaves • right contorted (in the tropics) or mycotrophs. • glabrous - no hairs!

Gentiana Symbolanthus

Voyria

Blackstonia Gentiana

2 *Gentianaceae - gentians *Gentianaceae - gentians CA (4-5) CO (4-5) A 4-5 G (2) Gentiana is 5 merous, with plaits between each lobe • flowers 4 or 5 merous Gentiana • pistil superior of 2 carpels • parietal placentation; capsular

Gentiana andrewsii - Gentiana puberulenta - Bottle gentian Prairie gentian

*Gentianaceae - gentians *Gentianaceae - gentians Gentianopsis is 4 merous, with fringed Gentianella is 4-5 merous, without petals fringe, and without plaits between petal lobes

Gentianopsis crinita - fringed gentian Gentianopsis procera - fringed gentian Gentianella quinquefolia - stiff gentian

3 *Gentianaceae - gentians *Apocynaceae - milkweeds • woody gentians common in cloud forests of the Worldwide family of trees, , herbs with opposite leaves - 415 genera, Neotropics 4600 species. • hummingbird, bat, and bee pollinated radiations

Symbolanthus Lisianthius

Pachypodium Periploca

*Apocynaceae - milkweeds *Apocynaceae - milkweeds Worldwide family of trees, vines, herbs with opposite leaves - 415 genera, Milky latex commonly poisonous and 4600 species. source of medicinal drugs.

Vinca minor - periwinkle Introduced ground cover, often spreading & monarch cardiac glycosides

Caralluma europea cactus mimic

Asclepias syriaca Catharanthus roseus common milkweed Malagasy periwinkle Curare - South America

4 *Apocynaceae - milkweeds *Apocynaceae - milkweeds

Family is broadly defined and includes distinctive milkweeds of old family CA (5) CO (5) A 5 G 2 primitive Asclepiadaceae • flowers 5 merous • left contorted perianth • 2 separate carpels - follicles Periploca

Transition to more elaborated ‘milkweed’ flowers Apocynum androsaemifolium Spreading dogbane

*Apocynaceae - milkweeds *Apocynaceae - milkweeds ______CA (5) CO (5) A 5 G 2 primitive CA (5) CO (5) A (5) G 2 derived

• ‘pollen presentation’ - style plunger or bottle brush to expose pollen • fuse to each other and to style region - (important! part of the suite of features in Asterid “heads” or pseudanthia) gynostegium • pollen forms pollinia • more with tufts of hairs

Asclepias - milkweed

• 5 stamens begin to be connivent

5 *Apocynaceae - milkweeds *Apocynaceae - milkweeds

crest

hood

Note 2 free carpels slightly fused at top Corona for reward

Corona = hood + crest

*Apocynaceae - milkweeds *Apocynaceae - milkweeds

Fusion of 5 stamens and top of Gland is attached to 2 pollinia

Corona = hood + crest Corona = hood + crest Gynostegium = A + G Gynoestegium = A + G Pollinia = pollen mass

6 *Apocynaceae - milkweeds *Apocynaceae - milkweeds

Gland is attached to 2 pollinia Bee removing pollinia or inserting pollinia into stigmatic cleft Corona = hood + crest Gynoestegium = A + G Pollinia = pollen mass

*Apocynaceae - milkweeds *Apocynaceae - milkweeds

“Las Vegas” strategy of pollination; but when occurs, all ∞ ovules are fertilized. Typically few follicles are produced per . Note seeds with coma attached for wind dispersal.

Apocynum androsaemifolium Apocynum cannabinum Asclepias amplexicaulis - Clasping milkweed Spreading dogbane Hemp dogbane

7 *Apocynaceae - milkweeds *Apocynaceae - milkweeds

Asclepias tuberosa - weed

Apocynum androsaemifolium Apocynum cannabinum Asclepias syriaca - Common Spreading dogbane Hemp dogbane milkweed Asclepias incarnata - Swamp milkweed

*Rubiaceae - coffee *Rubiaceae - coffee Cosmopolitan family, most diverse in tropics, of 550 genera and over Cosmopolitan family, most diverse in tropics, of 550 genera and over 10,000 species 10,000 species

• trees, shrubs, lianas, and herbs • opposite (whorled) leaves with inter-petiolar stipules • important drug family

Psychotria nervosa

Houstonia caerula- Cinchona - Galium (bedstraw) - azure bluets quinine are these stipules?

8 *Rubiaceae - coffee *Rubiaceae - coffee

CA (4-5) CO (4-5) A 4-5 G (2)

• 4 merous in temperate regions, 5 merous in tropics - tendency to cluster (pseudanthia!) • only epigynous family in Gentianales • fruit usually a 2-seeded drupe

Galium aparine - cleavers Galium - bedstraw Cleavers and bedstraws are numerous; separated Pentagonia Wisconsin by number of whorled leaves and fruit type Panama

*Rubiaceae - coffee *Rubiaceae - coffee

Mitchella repens Partridge berry, twin-berry

Note: paired flowers above Houstonia caerula- azure bluets with connate pistils; “twin” berry to right

9 *Rubiaceae - coffee *Rubiaceae - coffee

Cephalanthus occidentalis - buttonbush

Only shrub in Wisconsin; note “heads” of tiny 4 merous flowers “pseudanthia”

Isertia

Coffea arabica - coffee

*Rubiaceae - coffee *Rubiaceae - coffee

Psychotria nervosa Cephaelis tomentosa Psychotria nervosa Sally Kellerman hotlips Hot Lips Hoolihan • the giant Psychotria is paraphyletic and includes more specialized • the giant genus Psychotria is paraphyletic and includes more specialized genera (e.g., condensed, bracted inflorescences) “pseudanthia”! genera (e.g., condensed, bracted inflorescences)

10 *Rubiaceae - coffee Solanales

Hydnophytum formicarum • order within ‘lamiids’ or Asterid l group (in greenhouse #4) • 5 families and nearly 5,000 species dominated by Solanaceae (nightshade) and Convolvulaceae (morning glory) lamiids • no iridoids, alternate leaves, plicate corolla, calyx persistent in fruit

Solanaceae Convolvulaceae

The genus Hydnophytum - ant and campanulids epiphtyes - along with the whole subtribe Hydnophytinae - are now shown to be derived out of the genus Psychotria!

*Solanaceae - nightshades *Solanaceae - nightshades Large cosmopolitan family of herbs, shrubs, or trees with nasty compounds. CA (5) CO (5) A 5 G (2) Important for source of foods (potato, tomato, pepper) and drugs.

• 5- merous flowers • axile placentation • Alternate leaved • berry fruited • Cyme or dichasium inflorescence

Capsicum - pepper

Solanum carolinense - Solanum dulcamara Brugmansia Nicotiana - tobacco Horse nettle Bittersweet

11 *Solanaceae - nightshades *Solanaceae - nightshades CA (5) CO (5) A 5 G (2) CA (5) CO (5) A 5 G (2)

• stamens often terminal pored for buzz • often calyx persistent in fruit pollination by bees (remember Ericaceae?)

Solanum carolinense - Horse nettle Physalis alkekengi - Japanese lantern

*Solanaceae - nightshades *Solanaceae - nightshades

Solanum dulcamara Bittersweet

Solanum dulcamara - bitterweet Solanum carolinense - Horse nettle

12 *Solanaceae - nightshades *Solanaceae - nightshades

Physalis virginiana edible include . . . Ground cherry Note: calyx surrounding berry

Capsicum - pepper

Solanum - tomato & eggplant

*Solanaceae - nightshades *Solanaceae - nightshades drug plants include . . . drug plants include . . .

Nicotiana tabacum

Atropa belladona Atropa belladona Brugmansia Datura stramonium

13 *Solanaceae - nightshades *Solanaceae - nightshades

Datura stramonium Jimson-weed

Introduced, Datura stramonium Introduced, Datura stramonium hallucinogenic roadside Jimson-weed hallucinogenic roadside Jimson-weed weed weed . . . also called . . . also called thorn-apple thorn-apple

Wisconsin Medical Journal 2005

*Solanaceae - nightshades

Introduced, Datura stramonium hallucinogenic roadside Jimson-weed weed . . . also called thorn-apple

Wisconsin Medical Journal 2005

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