Download Systematic Implications of DNA variation in subfamily

Asterids – Part 1
“Basal” Asterids, Lamiids
Spring 2012
Fig. 8.1
Asterid characters
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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)
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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)
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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
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(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)
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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
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(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)
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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)
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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
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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)
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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)
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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