“Lower” Core Tricolpates: The Caryophyllales and Saxifragales

Spring 2011

“Lower”Core tricolpates

Figure 9.3 from the text

Basal and “lower” core eudicotsOrder Ranunculales

Ranunculaceae – ButtercupsPapaveraceae - Poppies

Order ProtealesPlatanaceae - Sycamore

Order CaryophyllalesCaryophyllaceae - CarnationsAmaranthaceae - Amaranths“Portulacaceae” - PurslanesCactaceae - Cacti

Polygonaceae - Knotweeds

Order SaxifragalesCrassulaceae - StonecropsSaxifragaceae - Saxifrages

Basaltricolpates

“Lower”coretricolpates

“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

BetalainPigments

(alkaloidal)

AnthocyaninPigments(phenolic)

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, 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; swollen nodes; notched “petals”

• Special uses: Many ornamentals• Required taxa: Dianthus (pinks, carnations), Silene

Caryophyllaceae: Dianthus

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

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

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 C4 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 pearareole; 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 eudicotsOrder Ranunculales

Ranunculaceae – ButtercupsPapaveraceae - Poppies

Order ProtealesPlatanaceae - Sycamore

Order CaryophyllalesCaryophyllaceae - CarnationsAmaranthaceae - Amaranths“Portulacaceae” - PurslanesCactaceae - Cacti

Polygonaceae - Knotweeds

Order SaxifragalesCrassulaceae - StonecropsSaxifragaceae - Saxifrages

Basaltricolpates

“Lower”coretricolpates

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

“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: Sedum

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

Sedum (Stonecrop)

-leaf blades terete or subterete, margins entire-inflorescences cymose-fruits follicles-largest genus in the family, likely to be split further

“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

• Family not required; for information only

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!