Ceratophyllales, “Basal” Eudicots,
Caryophyllales
Spring 2014
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 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 family
Ranunculaceae
Digital
Flowers
Ranunculus: buttercup
Ranunculaceae – Ranunculus
Buttercups
Ranunculaceae – Aquilegia
Columbines
Nectar spur
Spring Ephemerals
“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
“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
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 family
Polygonaceae - ocrea
Polygonaceae: Polygonum
(knotweeds)
-a number of species in this
genus are weedy
Polygonaceae:
Persicaria (smartweeds)
-a number of these are
native to North American
prairies, found especially in
potholes and sloughs
Polygonaceae Eriogonum
Polygonaceae – Buckwheat (Fagopyrum)
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
Core Caryophyllales
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
Required family
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 family
Amaranthaceae: Amaranthus
(amaranths)
Amaranthaceae: Chenopodium
(lamb’s quarters, quinoa)
Amaranthaceae: Salicornia (pickleweed)
-salt tolerant
-C4 photosynthesis
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 family
Cactaceae distribution
is restricted to the
western Hemisphere
except for Rhipsalis
Rhipsalis
Cactaceae
glochids
spine
Opuntia - Prickly pear
areole; glochids
(irritating hair-like
spines)
spines
Areole – axillary bud area
Cactaceae – Primitive genus Pereskia
-many with leaves
-spines but no
glochids
Cactaceae: Opuntia
-stem segments
flattened - “pads”
-glochids present
Cactaceae - Subfamily Cactoideae
-no leaves
-no glochids
Some cacti are bat pollinated!
https://www.youtube.com/watch?v=PJc1IhESV8c
https://www.youtube.com/watch?v=bZTbihSpMo8