1. “Lower” Core Tricolpates: The Caryophyllales and Saxifragales Spring 2010

2. “Lower” Core tricolpates Figure 9.3 from the text

3. Basal and “lower” core eudicots Order Ranunculales Ranunculaceae – Buttercups Berberidaceae - Barberries Papaveraceae - Poppies Order Proteales Platanaceae - Sycamore Order Caryophyllales Caryophyllaceae - Carnations Amaranthaceae - Amaranths “Portulacaceae” - Purslanes Cactaceae - Cacti Polygonaceae - Knotweeds Order Saxifragales Crassulaceae - Stonecrops Saxifragaceae - Saxifrages Hamamelidaceae - Witch hazels Basal tricolpates “Lower” core tricolpates

4. “Lower” Core Eudicots: The Caryophyllales Vessel elements with simple perforation plates Anther wall development Molecular data Two main clades: suborder Caryophyllineae and suborder Polygonineae (these sometimes treated as the orders Caryophyllales and Polygonales) 10,650 species in 30 families

5. Core of Suborder Caryophyllineae Repeatedly demonstrated to be monophyletic based on a range of 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 with ‘beaked’ integuments – Bitegmic ovules; inner set of integuments extend beyond outer set 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-element plastids (P-type)

6. Suborder Caryophyllineae Betalain pigments Anthocyanin pigments Figure 4.51 from the text

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

8. Caryophyllales - Caryophyllineae: Caryophyllaceae (The Carnation Family) Widespread, usually in temperate/warm temperate regions of N. hemisphere Herbs; leaves opposite, entire, sometimes hairy Diversity: Approx. 2,200 species in 70 genera Flowers: Tepals 4-5; carpels 2-5, superior ovary; stamens 4-10; fruit usually a loculicidal capsule Significant features: Presence of anthocyanin pigments; swollen nodes; notched petals Special uses: Many ornamentals Required taxa: Dianthus (carnations, pinks)

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

10. Caryophyllales - Caryophyllineae: Amaranthaceae (The Pigweed or Amaranth Family) Cosmopolitan, in disturbed, arid or saline habitats Primarily herbs, or small shrubs, occasionally succulent Diversity: Approx. 2,360 species in 169 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; many with C 4 photosynthesis Special uses: beets (Beta), spinach (Spinacia), amaranth (Amaranthus), and goosefoot (Chenopodium) are eaten as vegetables or pseudograins; ornamantals, agricultural weeds Required taxa: Amaranthus, Chenopodium

11. Amaranthaceae: Amaranthus (amaranths) -mainly annuals -leaves alternate, entire, petioles distinct -flowers small, unisexual, green or purplish -tepals 0 or 3-5, distinct, not withering when dry, chaffy

12. Amaranthaceae: Chenopodium (lamb’s quarters, quinoa) -annual or perennial -plants usually whitish (from hairs or glands) -leaves alternate, petiolate or sessile -flowers bisexual -tepals (3-) 5, usually basally connate, + soft

13. Caryophyllales - Caryophyllineae: “Portulacaceae” (The Purslane Family) Widely distributed in tropical and semi-tropical regions; diverse in W North America and Andes Succulent herbs Diversity: Approx. 450 species in 20 genera Flowers: Tepals 4-6, petal-like; stamens 4-6; carpels 2-3, in a superior or inferior ovary; 4 bracteoles associated with each flower, the inner 2 often calyx-like; fruit a loculicidal or circumcissile capsule (pyxis) Significant features: sometimes with CAM metabolism Special uses: Purslane (Portulaca oleracea) is eaten as a vegetable; many grown as ornamentals (Lewisia, Portulaca, Talinum) Required taxa: *family only

14. Caryophyllales - Caryophyllineae: 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, usually 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 taxa: Opuntia

15. Cactaceae Opuntia - Prickly pear areole; glochids! Areole – axillary bud area

16. Cactaceae – Primitive genus Pereskia

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

18. Cactaceae - Subfamily Cactoideae

19. Caryophyllales - Polygonineae: 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 usually of 6 petaloid tepals; stamens 5-9; carpels 2-3 in superior ovary; fruit an achene or nutlet, often 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 Special uses: buckwheat (Fagopyrum) fruits used as food; rhubarb (Rheum) petioles and sorrel (Rumex) leaves used as vegetable; many weeds Required taxa: Polygonum

20. Polygonaceae - ocrea

21. Polygonaceae: Polygonum (knotweeds) -herbs -ocrea often membranous -flowers bisexual -tepals 5

22. Basal and “lower” core eudicots Order Ranunculales Ranunculaceae – Buttercups Berberidaceae - Barberries Papaveraceae - Poppies Order Proteales Platanaceae - Sycamore Order Caryophyllales Caryophyllaceae - Carnations Amaranthaceae - Amaranths “Portulacaceae” - Purslanes Cactaceae - Cacti Polygonaceae - Knotweeds Order Saxifragales Crassulaceae - Stonecrops Saxifragaceae - Saxifrages Hamamelidaceae - Witch hazels Basal tricolpates “Lower” core tricolpates

23. Saxifragales Floral apex concave early in its development Carpels free (at least apically) Many also have a hypanthium Have retained 5-merous flowers with distinct parts Nearly 2,500 species in 14 families

24. “Lower Rosids” Saxifragales: Saxifragaceae (The Saxifrage family) Widely distributed in temperate and arctic regions, especially in N. Hemisphere Herbs Diversity: 550 species in 30 genera Flowers: with a variously developed hypanthium; sepals & petals usually 4 or 5, petals often clawed; carpels 2 (-5), + connate, ovary superior to inferior; fruit a septicidal capsule or follicle Significant features: stipules lacking; carpels usually fewer than the petals Required taxa: *family only *change from lab manual

25. “Lower” Rosids: Saxifragales: Crassulaceae (The Stonecrop Family) Widespread, from tropical to boreal regions; many from arid habitats Succulent herbs to shrubs, CAM metabolism Diversity: 1,500 species in 35 genera Flowers: Sepals & petals 4-5; carpels 4-5, free to slightly fused; a scale-like nectar gland subtends each carpel; fruit an aggregate of follicles Significant features: wide range of floral diversity; succulent leaves; lack a hypanthium; tiny seeds Special uses: ornamentals (Sedum, Sempervivum) Required taxa: Kalanchoe

26. Crassulaceae: Kalanchoe -mostly succulent herbs -flower parts in 4s or multiples (8 stamens) -petals connate, forming a tube -Old World; a few species commonly cultivated here

27. “Lower” Rosids: Saxifragales: Hamamelidaceae (The Witch Hazel Family) Scattered distribution in tropical and temperate regions Shrubs or trees with stellate hairs Diversity: 80 species in 25 genera Flowers: Sepals & petals usually 4-5; carpels 2, at least slightly connate, stigmas 2; fruit a loculicidal or septicidal woody capsule with woody exocarp & bony endocarp Significant features: anthers usually opening by 2 flaps Special uses: ornamental shrubs, witch hazel from extracts of bark of Hamamelis Required taxa: Hamamelis

28. Hamamelidaceae: Hamamelis -deciduous shrubs (rarely trees) -leaves alternate, oval, pinnately-veined -petals 4, long and narrow -stamens 4, alternating with 4 staminodia -capsules with explosive dehiscence, 1 seed per locule

29. Next time: On to the rosids!