Ceratophyllales , “Basal” Eudicots , Caryophyllales

1. Ceratophyllales, “Basal” Eudicots, Caryophyllales Spring 2014

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 “Basal” eudicots Order Ranunculales Ranunculaceae – Buttercups Berberidaceae - Barberries Papaveraceae - Poppies Order Proteales Platanaceae - Sycamore Order Caryophyllales Polygonaceae - Knotweeds Caryophyllaceae - Carnations Amaranthaceae - Amaranths Cactaceae - Cacti

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. 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 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

17. Papaveraceae - Papaver

18. “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

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) http://www.youtube.com/watch?v=O7eQKSf0LmY

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 Betalain Pigments Anthocyanin Pigments

37. CoreCaryophyllales 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 • Required family

39. Caryophyllaceae: Dianthus -”petals” 5-toothed, subtended by 2 or more overlapping bracts -stamens 10 -styles 2 -embryo scarcely curved

40. 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

41. 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

42. Amaranthaceae: Amaranthus(amaranths)

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

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

45. 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

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

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

48. Cactaceae – Primitive genus Pereskia -many with leaves -spines but no glochids

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

50. Cactaceae - Subfamily Cactoideae -no leaves -no glochids