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Asterids – Part 1 “Basal” Asterids, Lamiids Spring 2012 Fig. 8.1 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) • Iridoid compounds Asterid characters # stamens = # petals Unitegmic, tenuinucellate ovules Sympetaly and epipetalous stamens Fig. 8.83 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 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 “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 Davidia Cornaceae Nyssa (tupelo) 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 “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 Ericaceae anthers (Fig. 8.89) 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 Sarraceniaceae—another origin of carnivory by pitchers 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 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 petallike; 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 stylar 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 paired fruits anther views separate ovaries G: stylar head Apocynaceae – Groups without pollinia Vinca Plumeria Catharanthus Apocynum Thevetia Nerium oleander corona pair of pollinia Apocynaceae with pollinia (Asclepiadoideae) Ceropegia Hoya Calotropus Asclepias Stapelia 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 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 Rubiaceae interpetiolar stipules Rubiaceae Coffea arabica Pentas 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 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 solanacous tropane alkaloids (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 Solanaceae diversity 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 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) Fig. 8.83 Lamiales -gland-headed hairs -diacytic stomates -oligosaccharides (instead of starch) -anther anatomy -protein inclusions in the nuclei of mesophyll cells -endosperm with a micropylar haustorium -molecular data -ca. 22 families and 20,000 species 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 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 Lamiaceae From Zomlefer •Corolla: •zygomorphic •sympetalous •bilabiate •Stamens: •4, didynamous •Ovary: •epipetalous •2-carpellate •deeply 4-lobed •4 locules •Gynobasic style •Schizocarp (4 nutlets) Stachys floridana •Square stems (herbaceous taxa) •Opposite leaves •Inflorescence: false whorls (verticils or verticillasters) Lamiaceae Nepeta Monarda Phytostegia Note verticillate whorls of flowers at each node Ocimum Rosmarinus Clinopodium 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 Lamiaceae Economic plants and products: Condiments & perfumes: •Basil (Ocimum) •Lavender (Lavandula) •Oregano (Origanum) •Peppermint (Mentha) •Rosemary (Rosmarinus) •Sage (Salvia) •Spearmint (Mentha) •Thyme (Thymus) Mentha Lamiaceae Economic plants and products: Ornamental plants: •Beautyberry (Callicarpa) •Coleus (Coleus) •Salvia (Salvia) 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 Chionanthus Olea europaea Oleaceae Forsythia Syringa Oleaceae: Fraxinus -trees or shrubs -leaves usually pinnately compound -flowers small, usually unisexual -petals 4 or lacking -fruit a dry indehiscent samara Oleaceae: Syringa -upright shrubs -leaves simple -inflorescences paniculate -corolla salverform, lilac to pink or white -fruit a loculicidal capsule 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 Orobanchaceae diversity Agalinis Epifagus – beechdrops Conopholis Striga Pedicularis 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 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 Penstemon Plantaginaceae Digitalis Linaria Collinsia Antirrhinum Penstemon Plantaginaceae: Plantago -usually “stemless” herbs (rosettes) -leaves with parallel venation -flowers 4-merous, radial, windpollinated -corolla much reduced -stamens 4, filaments long exserted Plantaginaceae: Veronica -usually herbaceous, annuals or perennials -leaves usually opposite -flowers relatively small -corolla rotate, not bilabiate but somewhat zygomorphic -stamens 2 Lamiids: Lamiales: Scrophulariaceae (The Figwort Family) • • • • • • • Widely distributed in temperate to tropical regions Herbs (shrubs) with terminal inflorescences Diversity: 1,680 species in 52 genera Flowers: Sepals 3-5, connate; petals 4-5, connate, corolla bilabiate or tubular with flaring lobes, typically showing zygomorphy but sometimes actinomorphic; stamens, 5, 4, or 2, with filaments adnate to corolla; anther sacs usually confluent and opening by a single distal slit oriented at right angles to the filament or more or less Ushaped; carpels 2, connate, superior ovary; fruit a septicidal capsule, drupe or schizocarp of achenes or druplets Significant features: floral zygomorphy; anther openings Special uses: Many ornamentals (Buddleja, Nemesia, Scrophularia) Family not required Scrophulariaceae Scrophularia Leucophyllum Buddleja Sutera Scrophulariaceae: Verbascum -tall, usually hairy biennial herbs -stem leaves sessile, often decurrent -inflorescences terminal spikes, racemes or panicles -corolla 5-lobed, rotate, tube short, only slightly bilateral -stamens 5, all fertile, 3 or all of the filaments hairy Other cool Lamiales Bignoniaceae Lentibulariaceae – more carnivorous plants Gesneriaceae