Asterids – Part 1 Basal Asterids, Asterids I (Lamiids) Spring 2010

Phylogeny of the Major Angiosperm Groups RosidsAsterids Basal Tricolpates Caryophyllids Magnoliids & Monocots Basal Angiosperms Amborella Eudicots Core Eudicots tricolpate pollen

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

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)

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

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

“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

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 -calyx minutely 4-toothed -petals 4, stamens 4 -fruit a small drupe

“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

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

“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) Required taxa: Phlox

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

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

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

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

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

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

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

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) Required taxa: *family only *change from lab manual

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 *change from lab manual

Rubiaceae interpetiolar stipules

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

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 (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

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

Figure from the text

Apocynaceae with pollinia Asclepias Stapelia Ceropegia Hoya Calotropus

Figure from the text

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