1. Ceratophyllales, “Basal” Eudicots, Caryophyllales
Spring 2013

2. Major Angiosperm Clades
Amborellaceae
Nymphaeales
Austrobaileyales
MAGNOLIID
COMPLEX
MONOCOTS
EUDICOTS
[TRICOLPATES]
ANITA
GRADE
Soltis et al. 2000,
APG II 2002,
Judd et al. 2002

3. Fig. 7.1

4. 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

5. Ceratophyllum – Probably sister
to the eudicots
The most recent
molecular data
support this.

6. Ceratophyllales:
Ceratophyllaceae -
Ceratophyllum

7. Fig. 7.1

8. 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!

9. Fig. 8.1

10. “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

11. “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

12. Ranunculus: buttercup
Ranunculaceae
Digital
Flowers
Ranunculus: buttercup

13. Ranunculaceae – Ranunculus
Buttercups

14. Ranunculaceae – Aquilegia
Columbines
Nectar spur

15. Spring Ephemerals

16. “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 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

17. Papaveraceae - Papaver

18. “Basal” Eudicots: Proteales: Platanaceae (The Sycamore Family)
Tropical to temperate regions, N. America, S. Europe, SW & SE Asia
Trees
Diversity: 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

19. Platanaceae – Platanus occidentalis

20. Fig. 8.1

21. 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

22. Fig. 8.17

23. 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

24. Droseraceae – Drosera (sundews)

25. Dionaeaceae – Dionaea (Venus fly traps)

26. Nepenthaceae –
Nepenthes

27. 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

28. Polygonaceae - ocrea

29. Polygonaceae: Polygonum (knotweeds)
-a number of species in this
genus are weedy

30. Polygonaceae: Persicaria (smartweeds)
-a number of these are
native to North American
prairies, found especially in
potholes and sloughs

31. Polygonaceae -
Eriogonum

32. Polygonaceae – Buckwheat (Fagopyrum)

33. Fig. 8.17

34. 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

35. Core Caryophyllales
Fig. 8.18

36. Core Caryophyllales
Anthocyanin Pigments
Betalain Pigments

37. Suborder Caryophyllineae
Ovule and Seed Characters
Agrostemma sp.
curved embryo
perisperm
“Beaked” integument of ovule

38. 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

39. 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 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

40. Amaranthaceae: Amaranthus (amaranths)

41. Amaranthaceae: Chenopodium (lamb’s quarters, quinoa)

42. Amaranthaceae: Salicornia (pickleweed)
-salt tolerant
-C4 photosynthesis

43. 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

44. Cactaceae distribution is restricted to the western Hemisphere
except for Rhipsalis
Rhipsalis

45. Areole – axillary bud area
Cactaceae
Opuntia - Prickly pear
areole; glochids
(irritating hair-like
spines)
spines
Areole – axillary bud area

46. Cactaceae – Primitive genus Pereskia

47. Cactaceae: Opuntia
-stem segments
flattened - “pads”
-glochids present

48. Cactaceae - Subfamily Cactoideae

49. Some cacti are bat pollinated!