Ceratophyllales, “Basal” Eudicots, Caryophyllales

Spring 2013

Major Angiosperm Clades

Amborellaceae

Nymphaeales

Austrobaileyales

MAGNOLIID COMPLEX

MONOCOTS

EUDICOTS [TRICOLPATES]

ANITAGRADE

Soltis et al. 2000, APG II 2002, Judd et al. 2002

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

Order RanunculalesRanunculaceae – ButtercupsBerberidaceae - BarberriesPapaveraceae - Poppies

Order ProtealesPlatanaceae - Sycamore

Order CaryophyllalesPolygonaceae - KnotweedsCaryophyllaceae - CarnationsAmaranthaceae - AmaranthsCactaceae - Cacti

“Basal”eudicots

“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

Ranunculaceae

Ranunculus: buttercup

DigitalFlowers

Ranunculaceae – Ranunculus

Buttercups

Ranunculaceae – AquilegiaColumbines

Nectar spur

Spring Ephemerals

“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

“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

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)

• 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: Polygonum (knotweeds)

-a number of species in this genus are weedy

Polygonaceae: Persicaria (smartweeds)

-a number of these arenative to North Americanprairies, 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:

• 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 CaryophyllalesBetalain PigmentsAnthocyanin Pigments

Suborder CaryophyllineaeOvule and Seed Characters

curved embryo

“Beaked” integument of ovule

perisperm

Agrostemma sp.

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)

• 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(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 distributionis restricted to the western Hemisphere except for Rhipsalis

Rhipsalis

Cactaceae

Opuntia - Prickly pearareole; glochids(irritating hair-like spines)

Areole – axillary bud area

spines

Cactaceae – Primitive genus Pereskia

Cactaceae: Opuntia

-stem segments flattened - “pads”-glochids present

Cactaceae - Subfamily Cactoideae

https://www.youtube.com/watch?v=PJc1IhESV8c

https://www.youtube.com/watch?v=bZTbihSpMo8

Some cacti are bat pollinated!