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

“Lower” Core tricolpates Figure 9.3 from the text

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

“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

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)

Suborder Caryophyllineae Betalain pigments Anthocyanin pigments Figure 4.51 from the text

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

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)

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

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

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

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

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

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

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

Cactaceae – Primitive genus Pereskia

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

Cactaceae - Subfamily Cactoideae

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

Polygonaceae - ocrea

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

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

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

“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

“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

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

“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

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

Next time: On to the rosids!