ceratophyllales, “basal” eudicots, caryophyllales spring 2014
TRANSCRIPT
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Ceratophyllales, “Basal” Eudicots, Caryophyllales
Spring 2014
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Major Angiosperm Clades
Amborellaceae
Nymphaeales
Austrobaileyales
MAGNOLIID COMPLEX
MONOCOTS
EUDICOTS [TRICOLPATES]
ANITA
GRADE
Soltis et al. 2000, APG II 2002, Judd et al. 2002
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Fig. 7.1
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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
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Ceratophyllum – Probably sister to the eudicots
The most recent molecular data support this.
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Ceratophyllales:Ceratophyllaceae -Ceratophyllum
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Fig. 7.1
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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!
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Fig. 8.1
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“Basal” eudicots, Saxifragales, Vitales, Caryophyllales
Order RanunculalesRanunculaceae – ButtercupsBerberidaceae - BarberriesPapaveraceae - Poppies
Order ProtealesPlatanaceae - Sycamore
Order CaryophyllalesPolygonaceae - KnotweedsCaryophyllaceae - CarnationsAmaranthaceae - AmaranthsCactaceae - Cacti
“Basal”eudicots
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“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
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Ranunculaceae
Ranunculus: buttercup
DigitalFlowers
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Ranunculaceae – Ranunculus
Buttercups
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Ranunculaceae – AquilegiaColumbines
Nectar spur
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Spring Ephemerals
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“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
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Papaveraceae - Papaver
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“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
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Platanaceae – Platanus occidentalis
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Fig. 8.1
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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
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Fig. 8.17
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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
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Droseraceae – Drosera(sundews)
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Dionaeaceae – Dionaea (Venus fly traps)
http://www.youtube.com/watch?v=O7eQKSf0LmY
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Nepenthaceae – Nepenthes
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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
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Polygonaceae - ocrea
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Polygonaceae: Polygonum (knotweeds)
-a number of species in this genus are weedy
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Polygonaceae: Persicaria (smartweeds)
-a number of these arenative to North Americanprairies, found especially in potholes and sloughs
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Polygonaceae -Eriogonum
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Polygonaceae – Buckwheat (Fagopyrum)
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Fig. 8.17
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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
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Core Caryophyllales
Fig. 8.18
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Core Caryophyllales
Betalain PigmentsAnthocyanin Pigments
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Core Caryophyllales
Ovule and Seed Characters
curved embryo
“Beaked” integument of ovule
perisperm
Agrostemma sp.
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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
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Caryophyllaceae: Dianthus
-”petals” 5-toothed, subtended by 2 or more overlapping bracts-stamens 10-styles 2-embryo scarcely curved
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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
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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
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Amaranthaceae: Amaranthus(amaranths)
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Amaranthaceae: Chenopodium(lamb’s quarters, quinoa)
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Amaranthaceae: Salicornia (pickleweed)
-salt tolerant-C4 photosynthesis
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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
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Cactaceae distributionis restricted to the western Hemisphere except for Rhipsalis
Rhipsalis
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Cactaceae
Opuntia - Prickly pearareole; glochids(irritating hair-like spines)
Areole – axillary bud area
spines
spineglochids
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Cactaceae – Primitive genus Pereskia
-many with leaves-spines but no glochids
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Cactaceae: Opuntia
-stem segments flattened - “pads”-glochids present
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Cactaceae - Subfamily Cactoideae
-no leaves-no glochids
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https://www.youtube.com/watch?v=PJc1IhESV8c
https://www.youtube.com/watch?v=bZTbihSpMo8
Some cacti are bat pollinated!