Asterids – Part 1 Basal Asterids, Asterids I (Lamiids)

1. Asterids – Part 1 Basal Asterids, Asterids I (Lamiids) Spring 2011

2. Phylogeny of the Major Angiosperm Groups Eudicots Core Eudicots Basal Tricolpates Caryophyllids Rosids Asterids Magnoliids & Monocots Basal Angiosperms Amborella tricolpate pollen ANITA grade

3. Asterid characters • Molecular data • Ovules with a single integument • (reduction from two integuments) • Iridoid compounds

4. Core Asterids • Number of stamens = number of petals • Epipetalous stamens • Sympetalous corolla (also in Ericales) • Molecular data • Two main clades: lamiids (euasterids I: Garryales, Gentianales, Lamiales, Solanales) and campanulids (euasterids II: Aquifoliales, Apiales, Dipsacales, Asterales)

5. Asterid taxa “Basal” Asterids Order Cornales – dogwoods Order Ericales – azaleas, blueberries, cranberries Asterids I (lamiids) Order Solanales – potatoes, tomatoes, peppers Order Gentianales – gentians, milkweeds, coffee Order Lamiales – mints, olives, snapdragons Asterids II (campanulids) Order Apiales – ginseng, carrots, dill, parsley Order Dipsacales – honeysuckle, elderberry Order Asterales – bluebells, sunflowers Core Asterids

6. Asterid taxa – Part 1 “Basal” Asterids Order Cornales Cornaceae – dogwoods Order Ericales Ericaceae – blueberries, heaths Asterids I (lamiids) Order Solanales Solanaceae – potatoes, tomatoes, pepper Order Gentianales Rubiaceae – coffee, quinine Apocynaceae – dogbanes, milkweeds Order Lamiales Asterids II (campanulids)

7. “Basal” Asterids:Cornales: Cornaceae(The Dogwood Family) • Widespread, especially common in north temperate regions • Usually trees or shrubs; leaves usually opposite, usually entire, with secondary veins smoothly arching toward leaf margins (arcuate venation) • Diversity: 110 species in 7 genera • Flowers: Sepals & petals 4-5; stamens 4-10, pollen apertures with an H-shaped thin region; carpels usually 2 or 3, connate, inferior ovary; fruit a drupe, the pit winged or ridged • Significant features: Nectar disk on top of the ovary; inflorescences sometimes with showy bracts; basal lineage of the Asterids! • Special uses: Ornamentals such as (Cornus) and tupelo (Nyssa) • Required taxa: Cornus

8. Cornaceae: Cornus -shrubs, trees or herbs with usually opposite simple leaves -flowers small, in open cymes or in close heads surrounded by petal-like bracts (false flowers) -calyx minutely 4-toothed -petals 4, stamens 4 -fruit a small drupe

9. “Basal” Asterids:Ericales: Ericaceae(The Heath or Blueberry Family) • Cosmopolitan; most diverse in montane habitats in E. Asia, E North America, S Africa, Australia. Favor acid soils; sunny or part-shaded habitats • Trees, shrubs, lianas, occasionally mycoparasitic herbs lacking chlorophyll; leaves usually alternate and spiral • Diversity: 4,100 species in 124 genera • Flowers: Often showy. Sepals 4-5; petals 4-5, connate forming a cylindrical to urn-shaped corolla; stamens (3) 8-10; anthers often with appendages, and poricidal dehiscence, pollen grains often in tetrads; carpels 2-10, connate, superior to inferior ovary; fruit a septic. or loculic. capsule, berry, drupe • Significant features: anthers often with poricidal dehiscence & sometimes with appendages; leaves often coriaceous • Special uses: blueberries & cranberries (Vaccinium), Rhododendron and allies (Rhododendron, Erica, Kalmia, Pieris) are showy ornamentals • Required taxa: Rhododendron

10. Ericaceae: Rhododendron -shrubs or small trees with deciduous or evergreen leaves -flowers developed from scaly buds, mostly 5-merous -corolla deciduous -stamens usually 2x the number of corolla lobes; anthers with poricidal dehiscence -ovary superior -fruit a septicidal capsule

11. “Basal” Asterids:Ericales: Polemoniaceae(The Phlox Family) • Widely distributed; most diverse in temperate regions, especially western North America • Herbs, occasionally shrubs or small trees • Diversity: 380 species in 18 genera • Flowers: Sepals 5; petals usually 5, strongly connate forming a narrow tube, distal (free) ends of petals often markedly bent; stamens usually 5, filaments adnate to corolla tube; carpels 3, connate, superior ovary; fruit usually a loculicidal capsule • Significant features: Flowers distinctive with narrow tube and plicate and convolute corolla lobes • Special uses: Many ornamentals (Phlox, Gilia, Polemonium) • Family not required: for information only

12. Polemoniaceae: Phlox -herbaceous perennials (usually) -leaves usually opposite, simple, entire -flowers in cymes -stamens very unequally inserted on the long corolla tube -capsules ovoid

13. Asterid taxa – Part 1 “Basal” Asterids Order Cornales Cornaceae – dogwoods Order Ericales Ericaceae – blueberries, heaths Asterids I (lamiids) Order Solanales Solanaceae – potatoes, tomatoes, pepper Order Gentianales Rubiaceae – coffee, quinine Apocynaceae – dogbanes, milkweeds Order Lamiales Asterids II (campanulids)

14. Asterids I:Solanales: Solanaceae(The Potato Family) • Widespread but most diverse in the neotropics • Herbs, shrubs, trees, vines; leaves alternate; often with a ‘solanaceous smell’ • Diversity: 2,510 species in 102 genera • Flowers: Sepals 5, connate; petals 5, connate, forming variously tubular corolla, plicate (folded) ; stamens 5, filaments adnate to corolla, sometimes anthers connivant; carpels usually 2 (-5), connate, oriented obliquely to the median plane of the flower; superior ovary; fruit usually a berry (occ. a capsule, schizocarp or nutlet) • Significant features: Complex chemistry with solanacous tropane alkaloids; belladonna/atropine, nicotine, capsaicin, etc. • Special uses: Many useful fruits and vegetables (potatoes & tomatoes - Solanum, peppers - Capsicum), tobacco (Nicotiana), some ornamentals (Petunia) • Required taxa: Petunia, Solanum

15. Solanaceae: Solanum -herbs or shrubs -corolla regular, rotate, 5-merous, deeply lobed -anthers forming a tube around the style, with terminal openings; filaments short -fruit a berry, usually 2-locular -ca. 1,400 species, mostly tropical

16. Solanaceae: Petunia -herbs with upper leaves tending to become opposite -corolla slightly irregular, a little bilabiate, funnelform or salverform -stamens unequal, 1 much smaller than the others

17. Economic plants and products: Solanaceae • Edibles: • Cayenne pepper (Capsicum) • Eggplant (Solanum) • Green pepper (Capsicum) • Red pepper (Capsicum) • Potato (Solanum) • Tomato (Solanum)

18. Solanaceae Economic plants and products: • Medicinal/toxic plants • ~ Alkaloids! • Belladona (Atropa) • Henbane (Hyoscyamus) • Jimson-weed (Datura) • Nightshade (Solanum) • Mandrake (Mandragora) • Tobacco (Nicotiana)

19. Asterids I:Solanales: Convolvulaceae(The Morning Glory Family) • Widespread; most diverse in tropical, and subtropical regions • Twining and climbing herbs, often with laticifers; leaves alternate and spiral • Diversity: 1,930 species in 55 genera • Flowers: often showy; sepals 5, distinct or only slightly connate; petals 5, connate into funnelform tube, plicate and twisted in bud; stamens usually 5, filaments epipetalous, often of unequal lengths; carpels 2, connate, superior ovary; fruit a septifragal, circumcissile, or variously dehiscing capsule • Significant features: Latex (milky sap); flowers last for a day or less • Special uses: Sweet potato (Ipomoea batatas) root eaten as vegetable; some used as ornamentals, e.g. morning glory (Ipomoea); bindweed (Convolvulus), dodder (Cuscuta) • Family not required: for information only

20. Asterids I:Gentianales: Rubiaceae(The Coffee or Madder Family) • Cosmopolitan, most diverse in the tropics and subtropical regions • Trees, shrubs lianas or herbs, vines, shrubs; leaves opposite or whorled • Diversity: 9,000 species in 550 genera • Flowers: usually bisexual and radial; sepals 4-5, connate; petals 4-5, connate, forming a funnel shaped corolla; stamens usually 4 or 5, adnate to corolla; carpels usually 2 (-5), connate, inferior ovary; fruit a loculicidal or septicidal capsule, berry, drupe, or schizocarp • Significant features: interpetiolar stipules • Special uses: Major commodity is coffee (Coffea); anti-malarial drug obtained from the bark of Cinchona (quinine); ipecac (make-U-vomit) comes from Psychotria; gardenias (Gardenia), Pentas, and Ixora provide ornamentals • Required taxa: Galium

21. Rubiaceae interpetiolar stipules

22. Rubiaceae: Galium -stems 4-angled -slender herbs with whorled leaves -flowers small, in cymes -calyx teeth obsolete -corolla rotate -stamens 4 (rarely 3) -1 ovule per locule, the 2 carpels separating when ripe

23. Asterids I:Gentianales: Apocynaceae(The Milkweed Family; incl. Asclepiadaceae) • Widespread in tropical and subtropical regions; some in temperate regions • Trees, shrubs, herbs, lianas, vines with laticifers and usually milky sap • Diversity: 3,700 species in 355 genera • Flowers: Sepals usu. 5; petals usu. 5, connate forming bell- funnel- or tubular-shaped corolla; stamens usually 5, filaments always adnate to the corolla, anthers distinct or connate and forming a ring to fused to the stylar head; staminal outgrowths (corona) often present and petal-like; carpels usually 2, connate by styles/stigmas only & ovaries distinct to fully connate, superior ovary; apex of style expanded and highly modified, forming a 5-sided head, secreting viscin; fruits often paired, each ovary developing into a dry follicle, drupe or berry • Significant features: Usually opposite leaves; pollen in sticky masses (w/ viscin) or in pollinia; seeds flattened, often with a tuft of hairs • Special uses: Some chemical uses (e.g. Catharanthus, “Madagascar periwinkle”), and ornamentals (Asclepias, Vinca, Plumeria, Nerium) • Required taxa: Asclepias

24. Apocynaceae – Groups without pollinia Vinca Plumeria Catharanthus Apocynum Thevetia Nerium oleander

25. Figure 9.120 from the text

26. Apocynaceae with pollinia Ceropegia Hoya Calotropus Asclepias Stapelia

27. Figure 9.121 from the text

28. Apocynaceae: Asclepias -plants herbaceous, stems erect to leaning -leaves usually opposite, sometimes alternate or whorled -inflorescence an umbel -corona of 5 hooded fleshy bodies, each usually with an incurved horn but lacking a crest -pollen in pollinia, the pollinia suspended -fruit a dry, ovoid or lanceolate follicle, one of the pair often aborting