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Ceratophyllales , “Basal” Eudicots , Caryophyllales. Spring 2013. Major Angiosperm Clades. Amborellaceae Nymphaeales Austrobaileyales MAGNOLIID COMPLEX MONOCOTS EUDICOTS [TRICOLPATES]. ANITA GRADE. Soltis et al. 2000, APG II 2002, Judd et al. 2002. Fig. 7.1.

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Ceratophyllales , “Basal” Eudicots , Caryophyllales

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Ceratophyllales basal eudicots caryophyllales

Ceratophyllales, “Basal” Eudicots, Caryophyllales

Spring 2013

Ceratophyllales basal eudicots caryophyllales

Major Angiosperm Clades











Soltis et al. 2000,

APG II 2002,

Judd et al. 2002

Ceratophyllales basal eudicots caryophyllales

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

Ceratophyllales basal eudicots caryophyllales

Ceratophyllum – Probably sister

to the eudicots

The most recent

molecular data

support this.

Ceratophyllales basal eudicots caryophyllales


Ceratophyllaceae -


Ceratophyllales basal eudicots caryophyllales

Fig. 7.1

Eudicots tricolpates

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!

Ceratophyllales basal eudicots caryophyllales

Fig. 8.1

Basal eudicots saxifragales vitales caryophyllales

“Basal” eudicots, Saxifragales, Vitales, Caryophyllales



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

“Basal” Eudicots:Ranunculales: Ranunculaceae(The Buttercup Family)

  • 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





Ranunculus: buttercup

Ceratophyllales basal eudicots caryophyllales

Ranunculaceae – Ranunculus


Ceratophyllales basal eudicots caryophyllales

Ranunculaceae – Aquilegia


Nectar spur

Ceratophyllales basal eudicots caryophyllales

Spring Ephemerals

Basal eudicots ranunculales papaveraceae the poppy family

“Basal” Eudicots:Ranunculales: Papaveraceae (The Poppy Family)

  • 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

Papaveraceae - Papaver

Basal eudicots proteales platanaceae the sycamore family

“Basal” Eudicots:Proteales: Platanaceae (The Sycamore Family)

  • 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

Platanaceae – Platanus occidentalis

Ceratophyllales basal eudicots caryophyllales

Fig. 8.1

Core eudicots the caryophyllales

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

Ceratophyllales basal eudicots caryophyllales

Fig. 8.17

One origin of carnivory there is another in the asterids

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

Droseraceae – Drosera(sundews)

Dionaeaceae dionaea venus fly traps

Dionaeaceae – Dionaea (Venus fly traps)


Ceratophyllales basal eudicots caryophyllales

Nepenthaceae –


Non core caryophyllales polygonaceae the buckwheat or knotweed family

Non-core Caryophyllales:Polygonaceae(The Buckwheat or Knotweed Family)

  • 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 3- angled, 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 - ocrea

Polygonaceae polygonum knotweeds

Polygonaceae: Polygonum (knotweeds)

-a number of species in this

genus are weedy

Ceratophyllales basal eudicots caryophyllales

Polygonaceae: Persicaria (smartweeds)

-a number of these are

native to North American

prairies, found especially in

potholes and sloughs

Ceratophyllales basal eudicots caryophyllales

Polygonaceae -


Ceratophyllales basal eudicots caryophyllales

Polygonaceae – Buckwheat (Fagopyrum)

Ceratophyllales basal eudicots caryophyllales

Fig. 8.17

Core caryophyllales

Core Caryophyllales

Demonstrated to be monophyletic based mainly on DNA data, but most also share the following derived characters:

  • 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 caryophyllales1

Core Caryophyllales

Fig. 8.18

Core caryophyllales2

Core Caryophyllales

Betalain Pigments

Anthocyanin Pigments

Suborder caryophyllineae

Suborder Caryophyllineae

Ovule and Seed Characters

Agrostemma sp.

curved embryo


“Beaked” integument of ovule

Core caryophyllales caryophyllaceae the carnation family

Core Caryophyllales:Caryophyllaceae(The Carnation Family)

  • 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

Core caryophyllales amaranthaceae the pigweed or amaranth family

Core Caryophyllales:Amaranthaceae(The Pigweed or Amaranth Family)

  • 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: Amaranthus(amaranths)

Amaranthaceae chenopodium lamb s quarters quinoa

Amaranthaceae: Chenopodium(lamb’s quarters, quinoa)

Amaranthaceae salicornia pickleweed

Amaranthaceae: Salicornia (pickleweed)

-salt tolerant

-C4 photosynthesis

Core caryophyllales cactacaceae the cactus family

Core Caryophyllales:Cactacaceae(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

Ceratophyllales basal eudicots caryophyllales

Cactaceae distribution

is restricted to the

western Hemisphere

except for Rhipsalis




Opuntia - Prickly pear

areole; glochids

(irritating hair-like



Areole – axillary bud area

Cactaceae primitive genus pereskia

Cactaceae – Primitive genus Pereskia

Cactaceae opuntia

Cactaceae: Opuntia

-stem segments

flattened - “pads”

-glochids present

Cactaceae subfamily cactoideae

Cactaceae - Subfamily Cactoideae

Ceratophyllales basal eudicots caryophyllales

Some cacti are bat pollinated!



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