commelinids diversity and evolution of monocots€¦ · syagrus - lilliput palm of paraguay jubaea...

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Diversity and Evolution of Monocots . . . palms, spiderworts, bananas, and pineapples . . . Commelinids 4 main groups: • Acorales - sister to all monocots • Alismatids inc. Aroids - jack in the pulpit • Lilioids (lilies, orchids, yams) non-monophyletic petaloid • Commelinids Arecales – palms Commelinales – spiderwort Zingiberales –banana Poales pineapple grasses & sedges Commelinids • strongly monophyletic! • bound ferulic acid in cell walls (fluoresce under UV with ammonium hydroxide added) • this feature allowed placement of Dasypogonaceae • largest group of monocots ranging from palms to grasses Commelinids Dasypogonaceae 4 genera - W Australia

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Page 1: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

Diversity and Evolutionof Monocots

. . . palms, spiderworts, bananas, andpineapples . . .

Commelinids4 main groups:

• Acorales - sister to all monocots• Alismatids

– inc. Aroids - jack in the pulpit• Lilioids (lilies, orchids, yams)

– non-monophyletic– petaloid

• Commelinids– Arecales – palms– Commelinales – spiderwort– Zingiberales –banana– Poales

– pineapple– grasses & sedges

Commelinids

• strongly monophyletic!

• bound ferulic acid in cell walls(fluoresce under UV withammonium hydroxide added)

• this feature allowed placement ofDasypogonaceae

• largest group of monocots rangingfrom palms to grasses

Commelinids

Dasypogonaceae

4 genera - W Australia

Page 2: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

Commelinids• theme: reduction of flower, loss ofnectar, loss of zoophily, evolution ofbracts

pickeral weed

rapatead

grass

bromeliad

*Arecaceae - palms• the order has one family - alsocalled Palmae

• 190 genera and 2400 species oftrees and shrubs

• tropics, subtropics, deserts,Mediterranean biomes

*Arecaceae - palms• greatest center of diversity inMalay archipelago, thenAmazonia

• depauperate in Africa, butdiverse in Madagascar

Malaysia

Madagascar

*Arecaceae - palms• Rattan palms - a plant group that honors the WallaceBiogeographic Line

• Asian distribution with few species passing through Sulawesior New Guinea

Rattan palm & genericdistributions

Page 3: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

*Arecaceae - palms

Syagrus - lilliput palm of Paraguay Jubaea - Chilean wine palm

Great morphological diversity: in stature

*Arecaceae - palms

Lodoicea maldivica - Seychelles palm or double nut

Great morphological diversity: largest seed of seed plants

Lodoicea maldivica - Seychelles palm or double nut

This genus of 1 species endemic to the Seychelles has generatedinterest in having the largest seed, and in that the shape of theseed has suggested the devil's work or aphrodisiacal properties.

What is unusualabout how thisspecies was firstdiscovered?

*Arecaceae - palmsGreat morphological diversity: largest leaf

Raffia - rattan. . . and largest inflorescence

Corypha

Page 4: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

Roystonia

*Arecaceae - palmsVegetative characteristics

• “woody” stems via primarythickening meristem or diffusesecondary growth

• essentially hardened leaf bases

• single apical meristem:succeptible to frost

*Arecaceae - palmsVegetative characteristics

• palmate or pinnate compound,sheathing, plicate or folded

*Arecaceae - palmsFloral characteristics

• inflorescence surrounded byspathe - once allied with aroids

*Arecaceae - palmsFloral characteristics

• flowers unisexual or bisexual

CA 3 CO 3 A 3,6,∞ G 3 or (3)

Page 5: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

*Arecaceae - palmsFloral characteristics

• fruit a 1-seeded berry or drupe

Areca Betel palm

Cocos coconut

Nypa fruticanssalt marsh

Calamus radicalisHawaii

*Arecaceae - palmsClassification: 5 subfamilies

• Nypoideae and Calamoideae are first diverging

*Arecaceae - palmsImportant palms: food

Cocos nucifera -coconut

Phoenix - Date palm

*Arecaceae - palmsImportant palms: food

Page 6: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

Chyrtostachys -sealing wax palm

*Arecaceae - palmsImportant palms: oil, wax

Elaeis - oilpalm

Roystonea - Royal Palm Washingtonia inSanta Monica

*Arecaceae - palmsImportant palms: horticulture

Palm House at Kew RoyalBotanic Garden - largestglass house in the worldwith the largest glass houseplant - Chilean wine palm

*Arecaceae - palmsImportant palms: horticulture

Commelinids4 main groups:

• Acorales - sister to all monocots• Alismatids

– inc. Aroids - jack in the pulpit• Lilioids (lilies, orchids, yams)

– non-monophyletic– petaloid

• Commelinids– Arecales – palms– Commelinales – spiderwort– Zingiberales –banana– Poales

– pineapple– grasses & sedges

Page 7: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

Commelinids• theme: reduction of flower, loss ofnectar, loss of zoophily, evolution ofbracts

pickeral weed

rapatead

grass

bromeliad

Commelinales + Zingiberales

• 2 closely related tropical orders• primarily nectar bearing but with losses• bracted inflorescences

pickeral weed

nectar

spiderwort

pollen only

heliconia

nectar + bracts

Commelinaceae - spiderwort

Tradescantia ohiensis - spiderwort

Family of small herbs with succulentstems, stems jointed; leavessheathing. Family does not producenectar, but showy flowers for insectpollen gathering.

Commelina erecta - Erect dayflower

Commelinaceae - spiderwort

Tradescantia ohiensis - spiderwort

Inflorescence often bracted

Rhoeo - Moses in a cradle

Page 8: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

Commelinaceae - spiderwort

Tradescantia ohiensis - spiderwort

Flowers actinomorphic orzygomorphic

Commelina communis - day flower

CA 3 CO 3 A 6 G (3)

Commelinaceae - spiderwort• species rich in pantropics,especially Africa

• floral diversity is enormous

Pontederiaceae - pickerel weedAquatic family of emergents or floaters.Water hyacinth (Eichhornia) fromtropical America is invasive species insubtropical areas of the world.

Eichhornia crassipesWater hyacinthinvading Florida

Pickerel weed has glossy heart-shaped leaves,superficially like Sagittaria but without net venation.Flowers are in congested showy purple inflorescences.

Pontederiaceae - pickerel weed

Pontederia cordata - Pickerel weed

Page 9: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

Pontederia cordata - Pickerel weed

Flowers are showy, insectpollinated with nectar glands -previously placed in Liliales!

Pontederiaceae - pickerel weed Haemodoraceae - kangaroo paw

Anigozanthus - kangaroo paw

Small family with floral nectar,species radiations in Australia andSouth Africa

Zingiberales• strongly supported group of 8 tropicalfamilies

• rhizomatous monocots with showy,nectared, but highly bracted flowers

• 3 shared features: 1. Parallel-pinnate leaves, oftendistichous

Zingiberales• strongly supported group of 8 tropicalfamilies

• rhizomatous monocots with showy,nectared, but highly bracted flowers

• 3 shared features: 2. Bracted flowersand inflorescences

Page 10: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

Zingiberales• strongly supported group of 8 tropicalfamilies

• rhizomatous monocots with showy,nectared, but highly bracted flowers

• 3 shared features: 3. Inferior ovary

Zingiberales

DNA-based Zingiberales “rhizogram” by John Kress

• order fairly well known based on DNA and morphology

• show interesting trends in (1) fusion of perianth and (2) stamenloss and staminode development

Costus floral pattern

3 fused sepals3 separate petals5 fused sterile anthers(labellum)1 fertile anther

S S

S

P

P PA

AA

AAA

Poales I - showy flowers4 main groups:

• Acorales - sister to all monocots• Alismatids

– inc. Aroids - jack in the pulpit• Lilioids (lilies, orchids, yams)

– non-monophyletic– petaloid

• Commelinids– Arecales – palms– Commelinales – spiderwort– Zingiberales –banana– Poales

– pineapple– grasses & sedges

• showy flowers,insect or birdpollinated

• +/- reducedflowers, insect orwind pollinated

• reducedflowers, windpollinated

Poales I - showy flowers

Page 11: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

*Bromeliaceae - pineapples• mainly epiphytic, but terrestrial as well ininhospitable regions

*Bromeliaceae - pineapples• key adaptations: CAM photosynthesis, modifiedtrichomes or scales, “tank” formation

Scales (water & nutrientuptake)

Tank (water impounding)

*Bromeliaceae - pineapples• key adaptations: CAM photosynthesis, modifiedtrichomes or scales, “tank” formation

• scales very visible inSpanish moss

Tillandsia usneoides inSouth Carolina live oaks

*Bromeliaceae - pineapples• preadaptations to carnivory in Brocchinia andCatopsis

Amino acidsradioactivelylabeled beingincorporated intothe scales ofBrocchinia

Brocchinia Catopsis

Page 12: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

*Bromeliaceae - pineapples• inflorescence heavily bracted and often theattractant

*Bromeliaceae - pineapples CA 3 CO 3 A 6 G (3) or G (3)

__

• petals showy, but not the sepals

• 2 sets of 3 stamens• superior or inferior ovary, withtwisted styles

• berry or capsule

*Bromeliaceae - pineapples

1 species of Pitcairnia in west Africa -vicariance or dispersal?

• bromeliads are an American family: 2600 species, 56genera

guava

*Bromeliaceae - pineapples• pineapple not native to Hawaii - along with two otheringredients of Hawaiian Punch

passion fruit

Page 13: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

*Bromeliaceae - pineapples• classification traditionally had three subfamilies

tillandsioids pitcairnioids bromelioids

Incan ceremonial dance

*Bromeliaceae - pineapples• tillandsioids

Vriesea

Tillandsia usneoides and T. grandis

*Bromeliaceae - pineapples• bromelioids

Achmea Neoregelia

Ananas -pineapple

*Bromeliaceae - pineapples• pitcairnioids

Puya

Brocchinia

Navia

Page 14: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

*Bromeliaceae - pineapples• 2 subfamilies naturalbased on DNA

• pitcairnioids broadlyparaphyletic

• Brocchinia sister to restof family

• origin of family inGuayana Shield of SouthAmerica

*Bromeliaceae - pineapplesGuayana Highlands of southernVenezuela and adjacent areas ofBrazil and Colombia - the higherelevation “tepuis” are raindrenched and extremely nutrientpoor

*Bromeliaceae - pineapplesWhen did theAtlantic disjunctionoccur?

Pitcairnia saxicolaCosta Rica

Picairnia feliciana in west Africa

Bromeliaceae

Rapateaceae

DNA tree calibratedwith monocot fossils

Long distancedispersal to Africa!African speciesdivergence is 15-13mya

*Bromeliaceae - pineapplesWhen did theAtlantic disjunctionoccur?

Atlantic split

Page 15: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

*Bromeliaceae - pineapples

• African speciesoriginated from Andes!

Where did theAfrican speciescome from?

Rapateaceae - a tepui family• 16 genera and nearly 100 species from the Guayana Shield

Rapateaceae - a tepui family• most species are pollinated by pollen-gathering bees

• hummingbird pollination has evolved once in a clade of twogenera

Is the African Mascolocephalus a product of Atlanticvicariance with closest Guayana Shield relatives, or a

product of long distance dispersal?

Rapateaceae - a tepui family• most species in the Guayana Shield but one in west Africa

Page 16: Commelinids Diversity and Evolution of Monocots€¦ · Syagrus - lilliput palm of Paraguay Jubaea Chilean wine palm Great morphological diversity: in stature *Arecaceae - palms Lodoicea

Recent long distance dispersal to Africa!African species divergence is 8-6 mywhereas Atlantic separation is 80+ mya

Rapateaceae - a tepui family

Atlantic split