Ceratophyllales, “Basal” Eudicots, Caryophyllales Spring 2013

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

Fig. 7.1

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

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

Ceratophyllales: Ceratophyllaceae - Ceratophyllum

Fig. 7.1

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!

Fig. 8.1

“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

“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

Ranunculus: buttercup Ranunculaceae Digital Flowers Ranunculus: buttercup

Ranunculaceae – Ranunculus Buttercups

Ranunculaceae – Aquilegia Columbines Nectar spur

Spring Ephemerals

“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

Papaveraceae - Papaver

“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

Platanaceae – Platanus occidentalis

Fig. 8.1

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

Fig. 8.17

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

Droseraceae – Drosera (sundews)

Dionaeaceae – Dionaea (Venus fly traps) http://www.youtube.com/watch?v=O7eQKSf0LmY

Nepenthaceae – Nepenthes

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

Polygonaceae - ocrea

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

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

Polygonaceae - Eriogonum

Polygonaceae – Buckwheat (Fagopyrum)

Fig. 8.17

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

Core Caryophyllales Fig. 8.18

Core Caryophyllales Anthocyanin Pigments Betalain Pigments

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

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

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

Amaranthaceae: Amaranthus (amaranths)

Amaranthaceae: Chenopodium (lamb’s quarters, quinoa)

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

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

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

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

Cactaceae – Primitive genus Pereskia

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

Cactaceae - Subfamily Cactoideae

Some cacti are bat pollinated! https://www.youtube.com/watch?v=PJc1IhESV8c https://www.youtube.com/watch?v=bZTbihSpMo8