Ceratophyllales, “Basal” Eudicots,
Saxifragales, Caryophyllales
Spring 2012
Major Angiosperm Clades
Amborellaceae
Nymphaeales
ANITA
GRADE
Austrobaileyales
MAGNOLIID
COMPLEX
MONOCOTS
Soltis et al. 2000,
APG II 2002,
Judd et al. 2002
EUDICOTS
[TRICOLPATES]
Fig. 7.1
Ceratophyllales:
Ceratophyllaceae
-Submerged aquatic with many
adaptations for this habitat
-Fossil record extends back to the
early Cretaceous
-Phylogenetic position still uncertain,
but clearly part of the early radiation
of angiosperms above the ANITA
grade
Ceratophyllum – Probably sister
to the eudicots
The most recent
molecular data
support this.
Ceratophyllales:
Ceratophyllaceae Ceratophyllum
Fig. 7.1
Eudicots (tricolpates)
• Monophyletic: tricolpate pollen,
slender filaments in stamens*, and
loss of ethereal oils
• Ca. 125 million years old as a lineage
• Ca. 75% of angiosperm diversity (at
least 160,000 species)
• Flower parts in whorls, with whorls
alternating*
*also happened in monocots!
Fig. 8.1
“Basal” eudicots, Saxifragales, Vitales,
Caryophyllales
“Basal”
eudicots
Order Ranunculales
Ranunculaceae – Buttercups
Berberidaceae - Barberries
Papaveraceae - Poppies
Order Proteales
Platanaceae - Sycamore
Order Saxifragales
Saxifragaceae - Saxifrages
Crassulaceae - Stonecrops, sedum
Hamamelidaceae – Witch hazel
Order Vitales
Vitaceae - Grapes
Order Caryophyllales
Polygonaceae - Knotweeds
Caryophyllaceae - Carnations
Amaranthaceae - Amaranths
Cactaceae - Cacti
“Basal” Eudicots:
Ranunculales: Ranunculaceae
(The Buttercup Family)
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Widespread, but predominantly of temperate and
boreal regions
Herbs or less often shrubs or vines
Diversity: 2,300 species in 47 genera
Flowers: receptacle short to elongated, parts in
spirals; tepals 4 to many; stamens numerous; 5+
free carpels; fruit usually an aggregate of follicles
or achenes
Significant features: wide range of floral diversity
and pollination syndromes, toothed or lobed leaves
Special uses: primarily ornamentals
Required taxa: Ranunculus, Aquilegia
Ranunculaceae
Digital
Flowers
Ranunculus: buttercup
Ranunculaceae – Ranunculus
Buttercups
-cauline leaves alternate
-flowers usually yellow
(rarely white)
-petals (3-) 5 or more
-petals flat or concave,
often with a nectarproducing spot or pit
at the base
-fruit an aggregate of achenes
Ranunculaceae – Aquilegia
Columbines -herbs with 2-3ternately compound
leaves
-sepals 5, colored like
the petals
-petals 5, all alike, each
with a nectar spur
-fruit an aggregate of
follicles
-hummingbird, bee & hawkmoth
pollination
“Basal” Eudicots:
Ranunculales: Berberidaceae
(The Barberry Family)
• Widespread, in temperate regions of
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Northern Hemisphere & the Andes in South
America
Shrubs or herbs
Diversity: 700 species in 15 genera
Flowers: petals & sepals (4-) 6; stamens 4-6
(to numerous); unicarpellate, superior
ovary; fruit a berry, seeds often arillate
Significant features: Yellow wood (!).
Special uses: primarily ornamentals
Family not required
“Basal” Eudicots:
Ranunculales: Papaveraceae
(The Poppy Family)
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Widely distributed in temperate regions; N. Hemisphere,
South Africa
Herbs or soft wooded shrubs
Diversity: 780 species in 43-44 genera
Flowers: Sepals 2 (-3) & quickly deciduous; petals 4 (6);
carpels 2+, connate, superior ovary; fruit a capsule
(poricidal or slits)
Significant features: Leaves often highly dissected or
lobed; latex/laticifers present; most taxa are poisonous
Special uses: poppy (Papaver somniferum) source of
opiate alkaloids, ornamentals
Family not required
Papaveraceae - Papaver
-herbs with white latex
-flower buds nodding
-sepals 2, petals 4
-stigmas forming a flat,
4-20-rayed crown
-capsule with poricidal
dehiscence
Papaveraceae – Papaver somniferum – opium poppy
“Basal” Eudicots:
Proteales: Platanaceae
(The Sycamore Family)
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Tropical to temperate regions, N. America, S.
Europe, SW & SE Asia
Trees
Diversity: 8-10 species in 1 genus
Flowers: densely arranged in a raceme of globose
heads; flowers small, unisexual, inconspicuous,
wind-pollinated; fruits are aggregates of achenes
associated with hairs in dense, globose clusters
Significant features: characteristic bark; leaves
usually with palmate venation; axillary buds
covered by an enlarged petiole base
Special uses: ornamental trees, lumber
Family not required
Platanaceae – Platanus occidentalis
Fig. 8.1
Saxifragales
• Floral apex concave early in its
development
• Carpels free (at least apically)
• Many also have a hypanthium
• Have retained 5-merous flowers with
distinct parts
• Nearly 2,500 species in 14 families
Core Eudicots:
Saxifragales: Saxifragaceae
(The Saxifrage family)
• Widely distributed in temperate and arctic
regions, especially in N. Hemisphere
• Herbs
• Diversity: 600 species in 30 genera
• Flowers: with a variously developed hypanthium;
sepals & petals usually 4 or 5, petals often
clawed; carpels 2 (-5), + connate, ovary superior
to inferior; fruit a septicidal capsule or follicle
• Significant features: stipules lacking; carpels
usually fewer than the petals, gynoecium lobed
• Family not required
Clawed petals
Saxifragaceae
Core Eudicots:
Saxifragales: Crassulaceae
(The Stonecrop Family)
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Widespread, from tropical to boreal regions; many from
arid habitats
Succulent herbs to shrubs, CAM metabolism
Diversity: 1,500 species in 35 genera
Flowers: Sepals & petals 4-5; carpels 4-5, free to slightly
fused; a scale-like nectar gland subtends each carpel;
fruit an aggregate of follicles
Significant features: wide range of floral diversity;
succulent leaves; lack a hypanthium; tiny seeds
Special uses: ornamentals (Sedum, Sempervivum)
Family not required
Sedum (Stonecrop)
-leaf blades terete or subterete,
margins entire
-inflorescences cymose
-fruits follicles
-largest genus in the family,
likely to be split further
Core Eudicots:
Saxifragales: Hamamelidaceae
(The Witch Hazel Family)
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Scattered distribution in tropical and temperate
regions
Shrubs or trees with stellate hairs
Diversity: 90 species in 27-29 genera
Flowers: Sepals & petals usually 4-5; carpels 2, at
least slightly connate, stigmas 2; fruit a loculicidal
or septicidal woody capsule with woody exocarp &
bony endocarp
Significant features: anthers usually opening by 2
flaps
Special uses: ornamental shrubs, witch hazel from
extracts of bark of Hamamelis
Family not required
Hamamelidaceae: Hamamelis
-deciduous shrubs
(rarely trees)
-leaves alternate, oval,
pinnately-veined
-petals 4, long and narrow
-stamens 4, alternating with
4 staminodia
-capsules with explosive
dehiscence, 1 seed per locule
Fig. 8.1
Core Eudicots:
Vitales: Vitaceae
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(The Grape Family)
Widely distributed; maximal diversity in tropical
regions
Lianas (vines) with tendrils opposite the leaves
Diversity: 800 species in 14 genera
Flowers: Sepals 4-6, small; petals 4-6; stamens
opposite the petals; carpels 2, connate, superior
ovary; nectar disk well developed; fruit a berry
containing 4 seeds
Significant features: Tendrils are modified
inflorescences
Special uses: grapes (Vitis) form the basis of the
wine industry; also table grapes and raisins; some
ornamentals
Family not required
Vitaceae: Vitis
-bark without lenticels
-pith brown
-leaves simple, mostly
rounded/cordate
-inflorescence cymose,
compound
-flowers 5-merous
-petals separating at the
base and falling without
expanding as a cap
-seeds usually pyriform
Fig. 8.1
Core Eudicots:
The Caryophyllales
• Vessel elements with simple perforation
plates
• Anther wall development
• Support mainly from molecular data
• Two main clades: Core Caryophyllales and
the non-core Caryophyllales
• Evidence now supports placement sister
to the Asterids; previously near base of
core eudicots
• 10,650 species in 30 families
Fig. 8.17
One Origin of Carnivory
(there is another in the Asterids)
• One clade of the non-core Caryophyllales
evolved carnivory (lost in one of the
families)
• At least three mechanisms: snap-traps,
pitchers, sticky glands
Droseraceae – Drosera
(sundews)
Dionaeaceae – Dionaea (Venus fly traps)
http://www.youtube.com/watch?v=O7eQKSf0LmY
Nepenthaceae –
Nepenthes
Non-core Caryophyllales:
Polygonaceae
(The Buckwheat or Knotweed Family)
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Widely distributed, usually in temperate regions
Herbs, shrubs, trees, or vines
Diversity: Approx. 1,100 species in 43 genera
Flowers: Perianth of 4-6 petaloid (sepaloid) tepals; stamens 5-9;
carpels 2-3 in superior ovary; fruit an achene or nutlet, often 3angled, often associated with remaining perianth parts
Significant features: Presence of a sheathing stipule, the ocrea,
at stem nodes (lost in Eriogonum); nodes often swollen; leaves
usually alternate, simple and spirally arranged; flowers in
fascicles, these variously arranged in inflorescences
Special uses: buckwheat (Fagopyrum) fruits used as food;
rhubarb (Rheum) petioles and sorrel (Rumex) leaves used as
vegetable; many weeds
Required taxa: Polygonum, Persicaria
Polygonaceae - ocrea
Polygonaceae: Polygonum
(knotweeds)
-prostrate to erect herbs
to shrubs
-ocrea often membranous,
silvery, glabrous, apically
2-lobed, often disintegrating
completely or into fibers
-inflorescences spikelike
or flowers solitary
-flowers bisexual
-tepals 5, connate for
3-70% of length
-stamens 3-8
Polygonaceae:
Persicaria (smartweeds)
-usually erect herbs
-ocrea papery, usually
tan, brown or reddish,
never apically 2-lobed,
often tearing with age
-inflorescence spikelike,
paniclelike or capitate
-tepals 4 or 5, connate for
1/2-2/3 of length
-stamens 5-8
Fig. 8.17
Core Caryophyllales
Demonstrated to be monophyletic based mainly on DNA data, but
most also share the following derived characters:
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Betalain pigments – Nitrogen-containing (alkaloidal) red and
yellow pigments that replace the anthocyanin (phenolic)
pigments found in most other land plants
Presence of perisperm in seeds – specialized diploid tissue
derived from the megasporangium
Ovules campylotropous with ‘beaked’ integuments – inner
integument extends beyond outer at micropyle
Placentation free-central to basal
Coiled or folded embryos in seeds
Uniseriate perianth – single whorl of tepals
Stamens maturing centrifugally – Innermost anthers mature first,
progressively moving to outside of whorl
Special form of sieve tube plastids surrounded by proteinaceous
filaments
Core Caryophyllales
Fig. 8.18
Core Caryophyllales
Anthocyanin Pigments
Betalain Pigments
Suborder Caryophyllineae
Ovule and Seed Characters
Agrostemma sp.
curved embryo
perisperm
“Beaked” integument of ovule
Core Caryophyllales:
Caryophyllaceae
(The Carnation Family)
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Widespread, usually in temperate/warm temperate regions of
N. hemisphere
Herbs; leaves opposite, entire, sometimes hairy
Diversity: Approx. 2,400 species in 70 genera
Flowers: Tepals 4-5, usually appearing as sepals; outer whorl
of stamens often very petal-like and called “petals”; stamens
4-10; carpels 2-5, superior ovary; fruit usually a loculicidal
capsule
Significant features: Presence of anthocyanin pigments (loss
of betalains); swollen nodes; notched “petals”
Special uses: Many ornamentals
Family not required
Caryophyllaceae: Dianthus
-”petals” 5-toothed,
subtended by 2 or more
overlapping bracts
-stamens 10
-styles 2
-embryo scarcely curved
Caryophyllaceae: Silene
(Campion)
-stipules absent
-sepals connate, shorter
than the “petals”
-flower parts hypogynous
-styles 3 or 5
-fruit valves 3-5 or splitting
into 6-10 teeth
Core Caryophyllales:
Amaranthaceae
(The Pigweed or Amaranth Family)
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Cosmopolitan, in disturbed, arid or saline habitats
Primarily herbs, or small shrubs, occasionally succulent
Diversity: Approx. 2,000 species in 174-175 genera
Flowers: small, tepals usually 3-5; carpels 2-3, usually in
superior ovary; inflorescences compact; fruit an achene,
utricle, or circumcissile capsule (pyxis) usually associated
with persistent perianth parts
Significant features: Includes “Chenopodiaceae”; many
halophytes; polyporate pollen; stipules lacking; basal
placentation; many with C4 photosynthesis
Special uses: beets (Beta), spinach (Spinacia), amaranth
(Amaranthus), and goosefoot (Chenopodium) are eaten as
vegetables or pseudograins; ornamantals, agricultural weeds
Required taxa: Amaranthus, Chenopodium
Amaranthaceae: Amaranthus
(amaranths)
-mainly annuals
-leaves alternate, entire,
petioles distinct
-flowers small, unisexual,
green or purplish
-tepals 0 or 3-5, distinct,
not withering when dry,
chaffy
Amaranthaceae: Chenopodium
(lamb’s quarters, quinoa)
-annual or perennial
-plants usually whitish
(from hairs or glands)
-leaves alternate, petiolate
or sessile
-flowers bisexual
-tepals (3-) 5, usually basally
connate, + soft
Core Caryophyllales:
Cactacaceae
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(The Cactus Family)
North and South America; usually in arid zones or
seasonally dry regions; tropics to temperate regions
Spiny stem succulents; trees, shrubs, globular forms,
vines, epiphytes, geophytes
Diversity: 1,400 species in 97 genera
Flowers: Tepals numerous, often highly colored, spirally
arranged; stamens numerous; carpels 3 to many in an
inferior ovary; fruit a berry
Significant features: Lateral shoots reduced to areoles,
associated with a spine or spine cluster; reduced in
subfamily Opuntioideae to glochids; CAM metabolism
Special uses: Fruits (tunas) and stems (nopales) of
Opuntia and some other genera are eaten; many grown
as ornamentals.
Required taxa: Opuntia
Cactaceae
Opuntia - Prickly pear
areole; glochids!
Areole – axillary bud area
Cactaceae – Primitive genus Pereskia
Cactaceae: Opuntia
-stem segments
flattened - “pads”
-glochids present
Cactaceae - Subfamily Cactoideae