Understanding Basal Asterids: Cornales and Ericales Families

1. Asterids – Part 1 “Basal” Asterids, Lamiids Spring 2012

2. Fig. 8.1

3. Asterid characters • Molecular data • Sympetalous corollas • Epipetalous stamens • Number of stamens = number of petals • Ovules with a single integument and a thin nucellus (reduction from two integuments) • Iridoidcompounds

4. Asterid characters # stamens = # petals Unitegmic, tenuinucellate ovules Sympetaly and epipetalous stamens

5. Fig. 8.83

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

7. Asterid taxa – Part 1 “Basal” Asterids Order Cornales Cornaceae – dogwoods Order Ericales Ericaceae – blueberries, heaths Lamiids Order Gentianales *Apocynaceae – dogbanes, milkweeds Rubiaceae – coffee, quinine Order Solanales *Solanaceae– potatoes, tomatoes, pepper Order Lamiales Campanulids (Part 2) *family required for recognition

8. “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: 80-85 species in ca. 2 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 • Special uses: Ornamentals such as (Cornus) and tupelo (Nyssa) • Family not required

9. Davidia Cornaceae Nyssa (tupelo)

10. 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

11. “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 inverted, 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 • Family not required

12. Ericaceaeanthers (Fig. 8.89)

13. 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

14. Sarraceniaceae—another origin of carnivory by pitchers

15. Asterid taxa – Part 1 “Basal” Asterids Order Cornales Cornaceae – dogwoods Order Ericales Ericaceae – blueberries, heaths Lamiids Order Gentianales *Apocynaceae – dogbanes, milkweeds Rubiaceae – coffee, quinine Order Solanales *Solanaceae– potatoes, tomatoes, pepper Order Lamiales Campanulids (Part 2) *family required for recognition

16. Lamiids: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: ca. 4,600 species in ca. 400 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 stylarhead, 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

17. paired fruits anther views separate ovaries G: stylar head

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

19. corona pair of pollinia

20. Apocynaceaewith pollinia (Asclepiadoideae) Ceropegia Hoya Calotropus Asclepias Stapelia

21. 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

22. Lamiids: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: Ca. 12,000 species in ca. 600 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 (connate 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 • Family not required

23. Rubiaceae interpetiolar stipules

24. Rubiaceae Pentas Coffea arabica

25. Rubiaceae:Galium(bedstraw) -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

26. Lamiids:Solanales: Solanaceae(The Potato or Nightshade Family) • Widespread but most diverse in the neotropics • Herbs, shrubs, trees, vines; leaves alternate; often with a ‘solanaceous smell’ • Diversity: 2,450 species in 91-102 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 solanacoustropanealkaloids (belladonna/atropine, nicotine, capsaicin, etc.); stems with internal phloem • Special uses: Many fruits and vegetables (potatoes & tomatoes - Solanum, peppers - Capsicum), tobacco (Nicotiana), some ornamentals (Petunia) • Required taxa: Petunia, Solanum

27. Solanaceae diversity

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

29. 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

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

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

32. Fig. 8.83

33. Lamiales -gland-headed hairs -diacyticstomates -oligosaccharides (instead of starch) -anther anatomy -protein inclusions in the nuclei of mesophyll cells -endosperm with a micropylarhaustorium -molecular data -ca. 22 families and 20,000 species

34. Major Families of Lamiales • *Lamiaceae – mints • Oleaceae – olives, ashes, lilacs • Orobanchaceae – louseworts, beechdrops, Indian paintbrushes • Plantaginaceae – snapdragons, vervains, penstemons • Scrophulariaceae – mulleins, figworts *family required for recognition

35. Lamiids:Lamiales: Lamiaceae(The Mint Family; Labiatae) • Cosmopolitan • Herbs, shrubs, trees; stems square in herbaceous taxa • Diversity: Ca. 6,800 species in 236-238 genera • Flowers: Sepals 5, connate, calyx radial or bilateral; petals 5, connate, bilabiate; stamens 4, didynamous to more or less equal; carpels 2, 2 ovules per carpel, connate, styles terminal to an often gynobasic, superior, often deeply 4-lobed ovary; fruit a drupe w/ 1-4 pits, an indehiscent 4-seeded pod, or a schizocarp splitting into 4 nutlets or drupelets • Significant features: Opposite leaves (usually); aromatic volatile compounds - mint oils; inflorescences with main axis indeterminate and determinate (cymose) lateral axes, these often congested into pseudowhorls (verticillasters) • Special uses: Many herbs: oregano (Origanum), basil (Basilicum), peppermint/spearmint (Mentha), sage (Salvia), thyme (Thymus); teak wood (Tectona); ornamentals (e.g., Salvia, Callicarpa) • Required taxa: Salvia

36. From Zomlefer Lamiaceae • Corolla: • zygomorphic • sympetalous • bilabiate • Stamens: • 4, didynamous • epipetalous • Ovary: • 2-carpellate • deeply 4-lobed • 4 locules • Gynobasic style • Schizocarp (4 nutlets) • Square stems (herbaceous taxa) • Opposite leaves • Inflorescence: false whorls (verticils or verticillasters) Stachys floridana

37. Lamiaceae Note verticillate whorls of flowers at each node Nepeta Phytostegia Ocimum Rosmarinus Monarda Clinopodium

38. Lamiaceae: Salvia -calyx bilabiate, its lower lip 2-lobed -corolla strongly bilabiate -anther-bearing stamens 2, ascending and parallel, the upper pair lacking or rudimentary -anther with an elongate filament-like connective articulated with the filament -ovary deeply 4-parted

39. Lamiaceae Economic plants and products: • Condiments & perfumes: • Basil (Ocimum) • Lavender (Lavandula) • Oregano (Origanum) • Peppermint (Mentha) • Rosemary (Rosmarinus) • Sage (Salvia) • Spearmint (Mentha) • Thyme (Thymus) Mentha

40. Lamiaceae Economic plants and products: • Ornamental plants: • Beautyberry (Callicarpa) • Coleus (Coleus) • Salvia (Salvia)

41. Lamiids:Lamiales: Oleaceae(The Olive Family) • Widely distributed in tropical and temperate regions. • Trees, shrubs (lianas) with usually opposite leaves • Diversity: Ca. 700 species in 24 genera • Flowers: Often small but can be showy; sepals & petals 4, connate; stamens 2 (-4), filaments adnate to corolla; carpels 2, connate, superior ovary; fruit a loculicidal or circumcissile capsule, berry, drupe • Significant features: Tetramerous flowers, peltate secretory trichomes • Special uses: Olives (Olea) used for fruits and oil; jasmine (Jasminum) used for flavoring and perfumery; ornamentals: lilac (Syringa), ash (Fraxinus), privet (Ligustrum), Forsythia • Family not required

42. Chionanthus Oleaeuropaea Oleaceae Syringa Forsythia

43. Oleaceae: Fraxinus -trees or shrubs -leaves usually pinnately compound -flowers small, usually unisexual -petals 4 or lacking -fruit a dry indehiscent samara

44. Oleaceae: Syringa -upright shrubs -leaves simple -inflorescences paniculate -corolla salverform, lilac to pink or white -fruit a loculicidal capsule

45. Lamiids:Lamiales: Orobanchaceae(The Broomrape Family) • Nearly cosmopolitan • Herbs, hemiparasitic or holoparasitic (lacking chlorophyll), often black or dark colored when dry • Diversity: Ca. 2100 species in 96-99 genera • Flowers: Sepals 5, connate; petals 5, connate, the corolla 2-lipped; stamens 4, didynamous, adnate to the corolla; carpels 2, connate, superior ovary; fruit a loculicidal or septicidal capsule • Significant features: Haustorial connections to terrestrial plant roots • Family not required

46. Orobanchaceae diversity Agalinis Epifagus – beechdrops Pedicularis Striga Conopholis

47. Orobanchaceae: Castilleja -hemiparasitic herbs with alternate leaves -inflorescences with colorful, showy bracts more conspicuous than the flowers themselves -calyx tubular -anther locules unequal and separated

48. Lamiids:Lamiales: Plantaginaceae(The Snapdragon Family) • Nearly cosmopolitan, but most diverse in temperate areas • Herbs or less commonly shrubs, often with terminal inflorescences; leaves alternate or opposite • Diversity: Ca. 1,820 species in 101 genera • Flowers: Usually bisexual and bilateral, but + radial in Plantago; sepals 4-5, connate; petals 5 (or appearing to be 4 due to fusion of 2 upper lobes), connate, usually with a 2-lipped corolla; stamens usually 4 (2), with filaments adnate to corolla; carpels 2, connate, superior ovary; fruit a septicidal capsule or schizocarp of 2 achenes • Significant features: floral zygomorphy; special type of glandular hair • Special uses: Many ornamentals (Antirrhinum, Penstemon, Veronica, Linaria); medical foxglove (Digitalis) • Family not required

49. Digitalis Penstemon Plantaginaceae Linaria Collinsia Penstemon Antirrhinum

50. Plantaginaceae: Plantago -usually “stemless” herbs (rosettes) -leaves with parallel venation -flowers 4-merous, radial, wind- pollinated -corolla much reduced -stamens 4, filaments long exserted