A Tour of The Flowering Plants

A Tour of the Flowering Plants
An Introduction to Angiosperm
Phylogeny Classification
With emphasis on Colorado natives
Priscilla Spears November 2007
Spears. 2007.Tour of Flowering Plants

Why make a new classification?
• To create a “natural” classification, one
that mirrors evolutionary history
• To have taxonomic groups that hold an
ancestor, all of its descendants and only
its descendants
• As a part of “Assembling the Tree of Life”,
an effort to place all organisms in a
branching classification that reflects
evolution

What is phylogeny?
• The roots are phylo-, “tribe” or “lineage”
• A related Greek word is a verb “to bring
forth”
• -geno, “birth, production” as in genesis.
• Think family trees and genealogy–
branching diagrams of evolutionary
(ancestor-descendent) relationships.

What data are used?
• Morphological data are not discarded, but
emphasis is on shared, derived features.
• Developmental data
• Biochemical data
• DNA data is often the key.
DNA is the “second fossil record”.
DNA technology is necessary.
Information technology continues to evolve.

Are these two plants related?

Of the water lily, the lotus, or the
sycamore tree, which two are
most closely related,?

Oxalidales
Fabales
Rosales
nitr Apiales Asterales
oge Malpighiales
Cucurbitales n-fi
xer
s
Aquifoliales
Dipsacales
eurosids I
Fagales
Sapindales Gentianales
II
ids
Myrtales Ericales
ster
eua
eurosids II sI Lamiales
euasterid
A “Map” of Malvales
Solanales
Brassicales Boraginaceae
angiosperm Geraniales Cornales
s
erid
ros
orders shows the
ast
ids
Vitales
Caryophyllales
branching Saxifragales
pattern and Santalales
relationships Ranunculales
Core
eudicots
Proteales Liliales
Alismatales Asparagales
Laurales
ids
Basal lilio Poales
nids
eudicots commeli
Commelinales
al
Magnoliales Eudicots bas
ts
Magnoliid co
s ono Arecales
Zingiberales
M Acorales
Piperales
Nymphaeales Austrobaileyales
Amborellales
Tree diagram © 2006 MBG Press
Ancestral Seed
Spears. 2007.Tour of Flowering Plants Plant
Basal Angiosperms
• Three small basal lineages
– Amborellales (Amborella of New Caledonia)
– Nymphaeales, water lily, pond lily
– Austrobaileyales, Illicium (star anise),
Schisandra (star vine of SE United States)
• Magnoliids, a.k.a. eumagnoliids:
– Laurales, spice bush, avocado, cinnamon
– Magnoliales, magnolia, paw-paw, nutmeg
– Piperales, black pepper, kava, peperomia,
dutchman’s pipe, lizard’s tail
Magnoliids
Magnoliales
Laurales
Piperales
Piperales

Basal angiosperms in Colorado
• Nymphaeaceae –
Nuphar lutea, pondlily
or spatterdock
• No Austrobaileyales
• No magnoliids

Monocots
• They are clearly monophyletic (“one tribe”)
• Three main branches
– Basal monocots
• Acorales, Alismatales
– Lilioid (petaloid) monocots
• Asparagales, Liliales, Dioscoreales, Pandanales
– Commelinid monocots
• Arecales (palms), Commelinales, Poales,
Zingiberales

What’s new in monocots?
• Duckweeds are placed in Araceae
• Break up of the lily family
– Most monocots with six showy tepals were
placed in Liliaceae.
– The former lily family has been distributed
between two orders, Liliaceae and
Asparagales.
• Orchids are placed in Asparagales

Basal monocots
Alismatales
Acorales
Basal monocots in Colorado
• Acorales – Acoraceae
– Acorus calamus near Ft. Collins
• Alismatales – Alismataceae
– Several species of Alisma (water plantain) and
Sagittaria (arrowhead)
• Other Alismatales
– Juncaginaceae – Triglochin (arrowgrass)
– Hydrocharitaceae – Elodea (waterweed); Najas (water
nymph, introduced)
– Potamogetonaceae – Potamogeton (pondweed)

Lilioid or Petaloid monocots
• These names are less
formal.
• This group can also be
called the core
monocots.
• Orders are Liliales,
Asparagales,
Pandanales (screw
pines), and Dioscoreales
(tropical yam, Æ).

Liliales vs. Asparagales
Liliales:
• Nectaries on tepal or
filament base
• Tepals often spotted
• Seeds are light-colored &
have an outer epidermis
Asparagales:
• Nectaries on the septa of
the ovary
• Tepals not spotted
• Outer layer of seeds is
dark (in dry-fruited
species) or not present
Liliales
Colchicaceae, autumn
crocus & bellwort family
Melanthiaceae, wand lily
and trillium family
Alstroemeriaceae,
Chilean lily family
Liliaceae, lily and tulip
family
Smilacaceae
Liliales in Colorado
• Liliaceae • Melanthiaceae
Erythronium (avalanche lily) Veratrum (cornhusk lily)
Fritillaria (fritillary) Anticlea (wand lily, death
Lilium (wood lily) camas)
Lloydia (alp lily) Toxicoscordion (death
Streptopus (twisted stalk) camas)
[Zigadenus]
• Colchicaceae
Trillium
Prosartes [Disporum] (fairy
bells) • Smilacaceae
Smilax (carrionflower)

Colorado Liliales
1 1, 2, 4:
Liliaceae
3:
Colchicaceae
2
1. Lilium
philadelphicum,
4
wood lily
2. Lloydia serotina,
alp lily
3. Prosartes
[Disporum], fairy
bells
4. Streptopus,
3 twisted stalk

Families pictured: Asparagales
Iridaceae, iris family
Alliaceae, onion
family
Orchidaceae, orchid
family
Agavaceae, agave
and yucca family
Other families:
Agapanthaceae
Amaryllidaceae
Asparagaceae
Asphodelaceae
Hemerocallidaceae
Hyacinthaceae
Ruscaceae
Themidaceae
Asparagales in Colorado
• Agavaceae • Orchidaceae
Yucca, Leucocrinum about 11 genera,
(sandlily) including Calypso,
• Alliaceae Corallorhiza,
Platanthera,
Allium (onion) Spiranthes
• Iridaceae • Ruscaceae
Iris Nolina (sotol)
Sisyrinchium (blue-eyed Maianthemum (false
grass) Solomon’s seal, star
• Themidaceae Solomon’s seal)
Androstephium, Triteleia

Commelinid monocots
Orders represented: Arecales – palm
Poales – grass, sedge, bromeliad, rush
Commelinales – spiderwort, pickerel weed
Zingiberales – banana, ginger, canna

Poales in Colorado
• Cyperaceae, sedge – Carex, 11 more genera
• Juncaceae, rush – Juncus, Luzula
• Poaceae, grass – See Janet Wingate
• Sparganiaceae, bur reed - Sparganium
• Typhaceae, cattail - Typha

Commelinales in Colorado
• Commelinaceae:
Commelina (dayflower)
Tradescantia (spiderwort)
• Pontederiaceae:
Heteranthera (mud plantain)
Zosterella
• Non-Colorado family – Haemodoraceae,
Anigozanthos (kangaroo paw from Australia)
is used as florist’s flower.

The eudicots are the largest clade
• Eudicots are the largest branch of
angiosperms, with about 75% of the
named species.
• They are also known as the tricolpates,
because their pollen has three apertures
or is derived from an ancestor with three
apertures.
• Fossil pollen dates them to at least 120
mya.

Why is the name “eudicot”?
• Dicots (dicotyledons) previously included
the magnoliids and basal lineages.
• This made dicots polyphyletic (“many
lineages”) – it included plants that were
only distantly related.
• The term “eudicot” (“true dicot”) denotes a
monophyletic (“single lineage”) group.

Basal eudicots - Ranunculales
Berberidaceae: Berberis, Mahonia,
Podophyllum (mayapple), Nandina
Ranunculaceae: buttercup, columbine
Papaveraceae: poppy, fumitory,
bleeding heart

Other basal eudicots
Buxales:
Buxaceae –
Buxus, boxwood
Proteales: (Ç)
Proteaceae Pachysandra,
Platanaceae Japanese spurge
Nelumbonaceae

Basal eudicots in Colorado
Ranunculales is our only native order.
• Berberidaceae – Berberis (barberry),
Mahonia (Oregon grape)
• Papaveraceae – Argemone (thistle
poppy), Papaver, Corydalis, Fumaria
• Ranunculaceae – Aconitum, Actaea,
Anemone, Aquilegia, Clematis,
Delphinium, Ranunculus, Thalictrum +

Papaveraceae: Corydalis caseana
Wolf Creek pass, July 11, 2006

Colorado Ranunculaceae
1
2
3
1. Delphinium
2. Actaea
3. Aquilegia
4. Aconitum
5. Psychrophila
4 5
Core eudicots
• Five main branches
– Caryophyllales
– Santalales
– Saxifragales
– rosids including eurosids I (fabids) &
eurosids II (malvids)
– asterids including euasterids I (lamiids) &
euasterids II (campanulids)

Caryophyllales
This order has a wide variety of adaptations
– Growth in arid or salty soils – cacti, ice plants,
glassworts, salt cedar, jojoba, greasewood
– Growth in nitrogen-poor environments via
carnivory – sundews, Venus flytrap
• Many families have betalain pigments.
• Leaves are simple; margins are entire.
• Only a few are mycorrhizal.
Caryophyllales
Illustrated families:
Amaranthaceae:
amaranth
Caryophyllaceae:
moss campion
Polygonaceae:
knotweed
Droseraceae:
sundew
Others include:
Ice plant
Four o’clock
Jojoba Cactus
Statice
Tamarisk
Caryophyllales in Colorado
• Amaranthaceae (includes the former
Chenopodiaceae): Amaranthus, Atriplex
(saltbush), Bassia, Chenopodium, Salicornia ++
• Cactaceae – 6 genera
• Caryophyllaceae: Arenaria, Cerastium,
Dianthus, Lychnis, Silene, Stellaria ++
• Droseraceae: Drosera (sundew)
• Nyctaginaceae: Abronia (sand verbena),
Mirabilis (four o’clock), Oxybaphus (umbrellawort)

More Caryophyllales in Colorado
• Polygonaceae: Bistorta, Eriogonum (sulfur
flower), Fallopia, Persicaria, Polygonum
(knotweed), Rumex +
• Montiaceae: Claytonia (spring beauty),
Lewisia, Oreobroma (pygmy bitterroot), Montia
[formerly listed in Portulacaceae]
• Sarcobataceae: Sarcobatus (greasewood)
• Tamaricaceae: Tamarix (salt cedar)
• Frankeniaceae: Frankenia jamesii

Colorado Caryophyllales
Top row:
Montiaceae
Lower row:
Cactaceae,
Nyctaginaceae

Santalales
• Most species are hemiparasites – tap host
for water and minerals
• Includes both epiphytic species (stem
parasites) and those that are rooted in soil
near the host (root parasites)
• Western mistletoes have greatly reduced
flowers, but some mistletoes have showy
flowers.

Santales: Santalaceae
Mistletoe: Arceuthobium, shown here growing on ponderosa pine
Staminate flowers Pistillate flowers

Santalales in Colorado
• Santalaceae: Arceuthobium (dwarf
mistletoes of pines and Douglas fir);
Phoradendron (mistletoes of cedar,
cottonwoods)
• These were formerly placed in
Viscaceae.
• This family is still being sorted out.

Saxifragales
• This order includes families that diverged
quickly, so are very diverse.
• Succulents, aquatics, trees, shrubs, vines,
and herbaceous plants
• A number have partly fused carpels with
free styles and stigmas.

Saxifragales
Crassulaceae
Saxifragaceae
Paeoniaceae Grossulariaceae

More Saxifragales
Altingiaceae
Hamamelis
Hamamelidaceae
Spears. 2007.Tour of Flowering Plants Liquidambar – sweet gum

Saxifragales in Colorado
• Crassulaceae: Sedum (stonecrop),
Rhodiola (king’s crown, rose crown)
• Grossulariaceae: Ribes (gooseberries,
currants)
• Haloragaceae: Myriophyllum (water milfoil)
• Saxifragales: Saxifraga, Heuchera,
Lithophragma, Mitella +

Colorado Saxifragales
1 2 3
5
1. Colorado currant
2. Snowball
saxifrage
3. King’s crown
4. Alpine alumroot
4
5. Brook saxifrage
Rosids
• Distinct petals in most; numerous stamens
common
• Vitales (grape family) may be a sister group or
basal branch
• Geraniales and Myrtales are rosids, but their
relationship to other rosids is uncertain
• Eurosids I or fabids (squash, beans, alders,
oaks, roses, oxalis, willows)
• Eurosids II or malvids (mustards, mallows, &
maples)

Rosids, basal and unplaced
Myrtales - Onagraceae
Vitales – Vitaceae, grapes
Geraniales - Geraniaceae

Rosid groups in Colorado
• Vitales – Vitaceae: Parthenocissus (Virginia
creeper), Vitis (grape)
• Geraniales – Geraniaceae: Erodium, Geranium
• Myrtales
• Lythraceae: Lythrum (winged loosestrife)
• Onagraceae: Chamerion (fireweed), Epilobium
(willowherb), Gaura, Oenothera (evening
primrose) +

Eurosids I – Nitrogen fixers
• Nitrogen-fixing clade has four orders:
– Cucurbitales
– Fabales
– Fagales
– Rosales
• Each of these orders has at least one
genus in one family that forms nitrogen-
fixing root nodules
• NOT ALL families form these associations

Eurosids I – Nitrogen-fixing orders
Cucurbitales – squash family

Fabales – bean* family
Fagales – alder, oak, & walnut families
Rosales – rose, buckthorn, Russian olive,
elm, fig, & nettle families
*Underline indicates nitrogen-fixers
Eurosids I – Colorado
• Fagales – Fagaceae: Quercus (oak)
Betulaceae: Alnus (alder), Betula (birch),
Corylus (hazelnut)
• Cucurbitales, Cucurbitaceae: Cucurbita
(gourd), Echinocystis (spiny wild cucumber)
• Fabales – Fabaceae: Astragalus, Dalea,
Lathyrus, Lupinus, Mimosa, Oxytropis,
Thermopsis, Trifolium, Vicia ++
Polygalaceae: Polygala (milkwort)

Eurosids I, Rosales, in Colorado
• Cannabaceae: Celtis (hackberry), Humulus
(hops)
• Elaeagnaceae: Elaeagnus (Russian olive,
introduced), Shepherdia (buffaloberry)
• Rhamnaceae: Ceanothus (buckbrush)
• Rosaceae: Amelanchier, Cercocarpus,
Crataegus, Dryas, Fragaria, Geum, Holodiscus,
Potentilla, Purshia, Rosa, Rubus, Sorbus
• Urticaceae: Urtica (nettle)
Underline indicates nitrogen-fixers

Other eurosids I
• Oxalidales, the oxalis family
• Malpighiales is a new
grouping with many diverse
families:
Euphorbiaceae, spurge
Hypericaceae, St. John’s
wort
Linaceae, flax
Passifloraceae,
passionflower
Salicaceae, willow, aspen
Violaceae, violet

Malpighiales in Colorado
• Euphorbiaceae: Agaloma (snow-on-the-
mountain), Croton, Chamaesyce, Euphorbia
(spurge), Tragia +
• Hypericaceae: Hypericum (St. John’s wort)
• Linaceae: Adenolinum, Linum, Mesynium (flax)
• Salicaceae: Populus (cottonwood, aspen), Salix
(willow)
• Violaceae: Viola (violet), Hybanthus (green violet)

Colorado eurosids I
1 2 3
1. Ceanothus fendleri, buckbrush

2. Cercocarpus montanus, mountain
mahogany
3. Oxytropis lambertii, Lambert’s loco
4,5. Aspen pistillate and staminate
catkins
4 5
Eurosids II (malvids)
• Brassicales – mustard, cleome, papaya, &
nasturtium families
• Malvales - rock rose & mallow families
Malvaceae includes the former Tiliaceae as a subfamily;
several other former families are now its subfamilies.
• Sapinales – poison ivy, citrus, & soapberry
families
Maples and horse chestnuts are placed in the
soapberry family (Sapindaceae) in its
Hippocastanoideae subfamily

Eurosids II
Mustards, Mallows, Maples a.k.a. malvids
Brassicales – Brassicaceae
Malvales – Malvaceae
Sapindales – Sapindaceae, horse chestnut
and maple subfamily

Eurosids II in Colorado
Brassicales:
• Cleomaceae: Cleome (bee plant), Polanisia
(clammy weed)
• Limnanthaceae: Floerkea (false mermaid)
• Brassicaceae: at least 45 genera, many introduced
Alyssum, Boechera, Cardamine, Descurainia,
Draba, Erysimum, Lepidium, Lesquerella,
Noccaea, Physaria, Sisymbrium ++

More eurosids II in Colorado
Malvales – Malvaceae are herbaceous here
Natives: Callirhoe (poppy mallow), Malva
(cheeseweed), Sidalcea, and Sphaeralcea (globe
mallow)
Aliens include Abutilon, Alcea, Hibiscus
Sapindales – Rutaceae, the citrus family
Ptelea (hoptree, wafer ash)
Sapindaceae: Sapindus (soapberry), Acer (maple)
Anacardiaceae: Rhus (poison ivy, sumac)
[Toxicodendron]

Colorado eurosids II
1. Cleome serrulata, Rocky

Mountain beeplant
2-3. Rhus aromatica var. trilobata,
three-leaf skunkbush, in flower and
with fruit
4. Sphaeralcea, globe mallow

Colorado Brassicaceae
1. Physaria, double
bladder pod
2. Erysimum,
alpine wallflower
3-4. Stanleya
pinnata, prince’s
plume, a
selenium
1 2 indicator plant
3 4
Asterids
• Basal orders:
– Cornales (dogwood, hydrangea, blazing star)
– Ericales (impatiens, heather, phlox, primrose,
pitcher plant, camellia, ocotillo, pitcher plant)
• Euasterids I or lamiids
– Boraginaceae, Gentianales, Lamiales,
Solanales
• Euasterids II or campanulids
– Apiales, Aquifoliales, Asterales, Dipsacales

Basal asterids in Colorado
Cornales – Cornaceae: Cornus (bunchberry, dogwood)
Hydrangeaceae: Jamesia (waxflower), Philadelphus
(mock-orange), Fendlera (fendler bush)
Loasaceae: Mentzelia (blazing star)
Ericales – Ericaceae: Arctostaphylos (kinnikinnick),
Gaultheria, Kalmia, Vaccinium (blueberry, bilberry),
Monotropa (pinesap), Pterospora (pinedrops), Chimaphila
(pipsissewa), Moneses, Pyrola +
Polemoniaceae: Gilia, Ipomopsis, Phlox, Polemonium+
Primulaceae: Androsace, Dodecatheon, Lysimachia,
Primula +

Basal asterids in Colorado
Cornales
waxflower, blazing star
Ericales,
kinnikinnic, shooting star

More basal asterids in Colorado
2
1 3
1. Androsace
4 5
2. Polemonium
3. Primula
4,5. Phlox
1, 2, 4. on Mt.
Goliath
3. Wet meadow,
South Park
5. Aurora reservoir
Euasterids
• Morphological
features:
fused petals
same number of stamens
as petals
stamens attached to the
corolla tube
usually a two-carpellate
pistil

Euasterids I (lamiids)
• Many have opposite leaves with smooth
(entire) margins and superior ovaries.
• The former Hydrophyllaceae is now a part
of Boraginaceae.
• Apocynaceae (Gentiales) now contains
Asclepiaceae, the milkweed family.
• Several genera of Verbenaceae moved to
Lamiaceae. (Vitex, Tectona – teak,
Callicarpa, Clerodendron)

Euasterids I
Gentianales
Boraginaceae
Solanales Lamiales

Euasterids I in Colorado
Boraginaceae: Eritrichium (alpine forget-me-not),
Hackelia, Lithospermum, Mertensia, Cryptantha,
Hydrophyllum, Nama, Phaecelia +
Solanales – Solanaceae: Datura, Physalis, Solanum+
Convolvulaceae: Calystegia, Convolvulus, Ipomoea
(bush morning glory), Cuscuta (dodder)
Gentianales – Gentianaceae: Eustoma, Frasera,
Gentiana, Gentianella, Gentianopsis, Swertia
Rubiaceae: Galium (bedstraw)
Apocynaceae: Amsonia, Apocynum, Asclepias

More euasterids I in Colorado
Lamiales – Lamiaceae: Agastache (hyssop),
Dracocephalum, Mentha, Monarda (bee balm), Salvia,
Scutellaria (scullcap), Stachys, Teucrium ++
Lentibulariaceae: Urticularia (bladderwort)
Martyniaceae: Proboscidea (unicorn plant)
Oleaceae: Forestiera (privet), Mendora
Phrymaceae: Mimulus (monkeyflower)
Verbenaceae: Glandularia, Verbena (many non-native
genera moved to Lamiaceae)
Scrophulariaceae has changed as shown in
subsequent slides.
Euasterids I in Colorado
Boraginaceae
1. Eritrichum, alpine
forget-me-not,
2. Phacelia sericea,
purple fringe,
Lamiaceae
3. Teucrium,
germander
Solanaceae
4. Quincula, purple
ground cherry

What’s happened to the scrophs?
• Traditional Scrophulariaceae was polyphyletic
(contained many lineages)
• Changes include:
– Orobanchaceae includes parasitic and
hemiparasitic species
– Plantaginaceae includes : Penstemon,
Veronica, Digitalis, Linaria, and Plantago
– Mimulus is placed in Phrymaceae

Orobanchaceae
Genera include (Ç) Orobanche, (Ç) Castilleja, Pedicularis (Ç),

Cordylanthus, and Rhianthus

Plantaginaceae
Colorado native genera include: Bacopa,

Besseya, Chionophila, Collinsia, Limosella,
Penstemon, Plantago, Veronica
Veronicastrum. Others include: Antirrhinum,
Digitalis, Hebe, Linaria
1
Scrophulariaceae
4
Colorado native: Scrophularia (Æ)

Others include: Buddleja (1), Diascia
(2), Nemesia (3), Verbascum (4).

Euasterids II (campanulids)
• Most have alternate leaves (but Dipsacales have
opposite), serrate-dentate leaf margins, inferior
ovaries
• Aquifoliales – Aquifoliaceae, holly family
• Apiales – Apiaceae, Araliaceae, & Pittosporaceae
• Asterales – plunger pollination
Asteraceae, Campanulaceae, Menyanthaceae
• Dipsacales – Adoxaceae, Caprifoliaceae, Linnaeaceae
Dipsacaceae, & Valerianaceae

Orders of euasterids II
Aquifoliales - Aquifoliaceae
Dipsacales - Dipsacaceae
Apiales
Apiaceae
Dipsacales
Caprifoliaceae
Asterales – shown later
Euasterids II in Colorado
Apiales – Apiaceae: Aletes, Angelica, Cicuta, Conium,
Harbouria, Heracleum, Ligustrum, Lomatium, Musineon,
Oreoxis, Osmorhiza, Periderida, Sium ++
Araliaceae: Aralia (wild sarsapilla)
Dipsacales – Adoxaceae: Sambucus, Viburnum
Caprifoliaceae: Symphoricarpos (Lonicera introduced)
Linnaeaceae: Linnaea (twinflower)
Valerianaceae: Valeriana (valerian)
(Dipsacaceae: Dipsacus (teasel) introduced)

Asterales - Plunger pollination
1. Platycodon stamens adhere closely to the style and release their pollen
2. The style captures the pollen and elongates. The stamens fall away.
3. The lobes of the stigma open and are receptive to pollen

Asterales in Colorado
Campanulaceae – Campanula (harebell),
Lobelia, Triodanis
Menyanthaceae – Menyanthes (buckbean)
Asteraceae – split into subfamilies and
tribes as illustrated subsequently

Asteraceae

Thistle subfamily, thistle tribe
Disk flowers with
long styles and
deep corolla
lobes
Spirals of
phyllaries
artichoke
cornflower
thistle

Chicory subfamily, lettuce tribe
Ligulate flowers have
five points at the end
of the corolla.
No disk flowers
Fruits with umbrella-
like pappus
lettuce
chicory
salsify
hawkweed
skeleton weed
sow thistle

Colorado Asteraceae:
Carduoideae & Cichorioideae
Carduoideae, Cardueae – the thistle tribe
Cirsium, plus many invasive introductions –
Arctium, Carduus, Centaurea, Onopordum
Cichorioideae, Cichorieae – the lettuce tribe
or ligulate composites – 5 corolla points
Agoseris, Crepis, Hieracium, Lactuca,
Lygodesmia, Stephanomeria

Aster subfamily, anthemis tribe
Disk flowers, with or without ray flowers

Leaves are typically aromatic and finely divided
Scaly pappus, if there is one
Artemsia, Achillea, Matricaria, Chrysanthemum

Aster subfamily, aster tribe
Most have both ray and
disk flowers
Pappus is bristles
Easter daisy, rabbitbrush,
erigeron, horseweed, are
shown; also contains
gumweed, goldenrod

Colorado Asteraceae
Asteroideae: Anthemideae & Astereae
Anthemideae, the anthemis tribe
Achillea (yarrow) and Artemisia (sagewort)
[Weber & Wittman place sagebrush in Seriphidium]
Astereae, the aster tribe members

Aster (1 species), Chrysothamnus, Ericameria,
Conyza, Erigeron, Grindelia, Heterotheca,
Machaeranthera, Solidago, Townsendia

Aster subfamily, coreopsis tribe
Two dissimilar
rows of bracts in
the involucre
Pappus often
has barbed
awns
Coreopsis, Cosmos,
Dahlia, Bidens

Aster subfamily, gayfeather tribe
Disk flowers only
Long appendages
on the two style
branches
Eupatorium
Ageratum
Liatris

Colorado Asteraceae
Asteroideae: Coreopsideae & Eupatorieae
Coreopsideae, the coreopsis tribe
Bidens (beggar’s ticks), Coreopsis,
Cosmos
Eupatorieae, the gayfeather tribe
Ageratina, Brickellia, Liatris (gayfeather)

Aster subfamily, everlasting tribe
Disk flowers
only
Phyllaries
(bracts of the
involucre) are
often papery
Strawflower
Pearly
everlasting
Pussytoes
Edelweiss

Aster subfamily, sneezeweed tribe
Ray flower corollas are often wedge-
shaped
Anthers are not dark
Alpine sunflower, Gaillardia, Helenium

Colorado Asteraceae
Asteroideae: Gnaphalieae & Helenieae
Gnaphalieae, the everlasting tribe
Anaphalis (pearly everlasting), Antennaria
(pussytoes), Gnaphalium,
Pseudognaphalium
Helenieae, the sneezeweed tribe
Gaillardia, Helenium, Hymenoxys,
Psilostrophe, Tetraneuris

Aster subfamily, sunflower tribe
Anthers are often dark
Typically two or more rows of phyllaries
Pappus may be scales, awns, or
bristles
Ragweed, coneflower, sunflower,
black-eyed Susan, zinnia, cocklebur

Aster subfamily, senecio tribe
Single row of phyllaries, all the same length,

often fused
Pappus is fine white bristles
Very large tribe that is widely adapted
Senecio, coltsfoot, Ligularia, string-of-pearls

Colorado Asteraceae
Asteroideae: Heliantheae & Senecioneae
Heliantheae, the sunflower tribe
Ambrosia (ragweed), Balsamorhiza,
Berlandiera, Echinacea, Helianthella,
Helianthus, Iva, Ratibida, Rudbeckia,
Silphium, Wyethia, Xanthium, Zinnia +
Senecioneae, the senecio tribe
Packera, Petasites, Senecio, Tetradymia,
Ligularia

For more information, see:
Spears, Priscilla. 2006. A Tour of the Flowering
Plants: Based on the classification system of
the Angiosperm Phylogeny Group.
ISBN 1-930723-48-2
Available from the publisher, Missouri Botanical Garden
Press and from Amazon.com.
308 pages, over 800 color photos
Stevens, P. F. (2001 and onwards) Angiosperm
Phylogeny Website
http://www.mobot.org/MOBOT/research/APweb/

A Tour of The Flowering Plants - An Introduction To Angiosperm Phylogeny November 2007

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Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

pattern and Santalales

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Wolf Creek pass, July 11, 2006

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Staminate flowers Pistillate flowers

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants Liquidambar – sweet gum

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Vitales – Vitaceae, grapes

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Cucurbitales – squash family

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

1. Ceanothus fendleri, buckbrush

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

1. Cleome serrulata, Rocky

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants

Spears. 2007.Tour of Flowering Plants