• 11 families and nearly 26,000 - Australasia appears to be center of Diversity and Evolution diversity of lamiids • no iridoids, latex common, inferior gynoecium, presentation . . . asters, ragweeds, and

goldenrods . . . campanulids

inferior G

bellflower - chickory - Campanulaceae

*Asteraceae - composites *Asteraceae - composites

One of the most successful of all flowering families with One of the most successful of all families with over 1,500 genera and 23,000 species over 1,500 genera and 23,000 species

• composites found throughout the world but most characteristic of the grassland biomes • but also diverse in arctic to tropical and subtropical regions

1 *Asteraceae - composites Pseudanthia in the Asterids

Family has 4 specialized features important in this radiation:

1. Special Cornaceae Apiaceae Rubiaceae “head” - pseudanthia

2. Pollen presentation

3. Diverse secondary chemistry

4. Whole genome duplication Asteraceae Caprifoliaceae Adoxaceae

Pathway to Asteraceae Head? Pathway to Asteraceae Head?

Menyanthaceae Goodeniaceae Asteraceae

How did this happen morphologically?

Pozner et al. 2012 (Amer J Bot)

2 Pollination Syndromes Pollen Presentation

hummingbirds flies moths bees & wasps butterflies wind Cross pollination Self pollination on inner receptive by curling of surfaces stigmas

Anthers fused forming a Pollen pushed out by a Stigma makes contact with tube for pollen release style that acts as a plunger self pollen if necessary

Chemical Diversity Chemical Diversity

Packera Agoseris Tanacetum Helenium hoopesii – Sneezeweed, Owl’s-claw • polyacetylenes, sequiterpene lactones, terpenes, • sequiterpene lactones poisonous to alkaloids, latex sheep in the southern Rockies • “spewing” disease

3 Chemical Diversity Whole Genome Duplication

• all Asteraceae have genome doubling – not in other Asterales

• species radiation associated with the WGD shows a “time lag”

Artemisia annua – Sweet wormwood

• sequiterpene lactones used in anti- malarial treatment • Chinese herbal medicine

Schranz et al. 2012 – Ancient whole genome duplications, novelty and diversification: the WGD Radiation Lag-Time Model.

Asteraceae - composites Asteraceae - floral diversity __ The head or capitulum is a cluster CA X CO (5) A (5) G (2) of 1 or 2 distinct flower types. The family is also called “Compositae” referring to this clustering.

The head is surrounded by special called the involucre or phyllaries. • Calyx is reduced to a pappus of scales, awns, bristles, or absent The involucre is important in the • Corolla has 5 but classification and identification within the variously fused or zygomorphic family. • Anthers (only) fused in a ring

4 Asteraceae - floral diversity Asteraceae - floral diversity __ Disc floret True Ray Ligulate floret CA X CO (5) A (5) G (2)

The fruit is a one-seeded with the pappus serving as the fruit disperser (e.g., barbs for animal dispersal, hairs for wind dispersal) • the family exhibits a diverse array of floral types • these three corolla types are seen in Wisconsin

Asteraceae - floral diversity Asteraceae - floral diversity

1 1 2

Main floret types Main floret types

1. Disk or tubular florets are actinomorphic 1. Disk or tubular florets are actinomorphic

2. Ray florets are usually 3 long fused petals + 2 obsolete petals

5 Asteraceae - floral diversity Asteraceae - head diversity

1 3 These various types of florets combine 2 to form a number of different looking heads

Radiate head: disk florets in the center, ray florets along the edge Main floret types (these usually pistillate)

1. Disk or tubular florets are actinomorphic

2. Ray florets are usually 3 long fused petals + 2 obsolete petals Symphyotrichum - aster 3. Ligulate florets are 5 fused petals but split open

Asteraceae - head diversity Asteraceae - head diversity

These various types of florets combine These various types of florets combine to form a number of different looking to form a number of different looking heads heads

Radiate head: disk florets in the Radiate head: disk florets in the center, ray florets along the edge center, ray florets along the edge (these usually pistillate) (these usually pistillate)

Discoid head: only disk or tubular Discoid head: only disk or tubular florets comprise the entire head florets comprise the entire head

Ligulate head: only ligulate florets Liatris - blazing star comprise the entire head Cichorium - chickory

6 Asteraceae - phylogeny Asteraceae - phylogeny • early DNA based work surprisingly placed a small group of • early DNA based work surprisingly placed a small group of South America genera as sister to the rest of the family and South America genera as sister to the rest of the family and recognition of 3 subfamilies and many tribes recognition of 3 subfamilies and many tribes

47·5 million-year- old from Patagonia

Extant member of an early branching lineage of Asteraceae (Mutisioideae), lorentzii lanceolata

Asteraceae - phylogeny Asteraceae - phylogeny • later DNA based work questioned the monophyly of • family originated in South America subfamily Cichorioideae - classification still in progress

7 Asteraceae - tribes Asteraceae - tribes Tribe Barnadesiinae Tribe Mutisieae Small South American tribe with bilabiate (C) South American tribe usually with bilabiate or pseudolabiate (D) corolla - similar to some other families of Asterales flowers; characteristic of tepuis in northern South America

disk ray ligulate

Barnadesia lanceolata Stomochaeta Duidaea Cerro Autana

Asteraceae - tribes Asteraceae - tribes Tribe Lactuceae (Cichorieae) Tribe Lactuceae (Cichorieae) Only have ligulate heads, milky latex in vegetative parts, pappus of filaments

Dandelion, goat’s-beard, chickory, hawkweed

Cichorium intybus - chickory Eurasian weed - totally naturalized Taraxacum - dandelion and distributed widely even to Botany 400 plant collections

8 Asteraceae - tribes Asteraceae - tribes Tribe Lactuceae (Cichorieae) Tribe Cardueae (Cynareae)

• spiny phyllaries and Cirsium - often and stems thistles • heads never radiate

• petals white or Hieracium - cynanic colors (blues, hawkweeds purples)

• thistle, knapweed, burdock

Sonchus asper – sow thistle

Asteraceae - tribes Asteraceae - tribes

Tribe Cardueae (Cynareae) Tribe Vernonieae Alternate leaves Purple flowers generally

Hesperomannii Hawaii

Centaurea maculosa - Spotted knapweed Arctium - burdock Vernonia fasciculata - ironweed

9 Asteraceae - tribes Asteraceae - tribes

Tribe Inuleae (Gnaphalieae) usually covered with white hairs, Tribe Inuleae (Gnaphalieae) pappus of bristles or hairs

Pussy toes, cudweed, pearly everlasting

Antennaria plantaginifolia - pussy toes

Anaphalis margaritacea - pearly everlasting

Asteraceae - tribes Asteraceae - tribes Tribe Astereae Tribe Astereae Conspicuous rays in radiate head, pappus of plumose bristles; asters, goldenrods, fleabanes

Symphyotrichum pilosus – frost aster Symphyotrichum novae-angliae Solidago rigida - annuus – (Botany 400 plant collections) - New England aster stiff goldenrod daisy fleabane

10 Asteraceae - tribes Asteraceae - tribes Tribe Anthemideae Tribe Anthemideae

Aromatic plants, with dissected, alternate leaves; pappus of awns or scales or missing Ox-eye daisy, yarrow, wormwood, tansy, dog fennel

Achillea millefolium – yarrow (Botany 400 plant collections) Tanacetum bipinnatrum – Matricaria discoidea - pineapple weed Leucanthemum vulgare Lake Huron tansy Ox-eye daisy, chrysanthemum

Asteraceae - tribes Asteraceae - tribes Tribe Senecioneae Tribe Helenieae Alternate leaved, radiate heads, 3-lobed Alternate or basal leaved, phyllaries in 1 ray flowers always widest at tip row, capillary pappus; ragwort, Indian plantain

Packera aureus - golden ragwort Helenium autumnale – aristata– common sneezeweed common blanket-flower

11 Asteraceae - tribes Asteraceae - tribes Tribe Heliantheae Large tribe with radiate, multi-layered Tribe Heliantheae phyllaries, rays mostly yellow, tendency for opposite leaves Coneflower, sunflower, rosinweed, ox- eye, black-eyed Susan, prairie dock, coreopsis, Peruvian daisy

Ratibida pinnata - coneflower Helianthus annuus - sunflower Biden cernuus - bur marigold Echinacea pallida - purple coneflower

Asteraceae - tribes Asteraceae - tribes

Tribe Ambrosieae Tribe Eupatorieae (or Heliantheae) Ambrosia artemisiifolia (or Heliantheae) ragweed Opposite or Unisexual and wind whorled leaves pollinated, rays absent, not fused Discoid heads Ragweed, cocklebur Corolla never yellow

Male florets Joe-pye weed, boneset, snakeroot, Female florets blazing star Eutrochium maculatum Eupatorium perfoliatum Joe-pye weed Boneset

12