Commelinids Diversity and Evolution of Monocots · Lodoicea maldivica - Seychelles palm or double...

Diversity and EvolutionDiversity and Evolutionof Monocotsof Monocots

. . .. . . palms, spiderworts palms, spiderworts, bananas,, bananas, and andpineapples pineapples . . .. . .

CommelinidsCommelinids4 main groups:4 main groups:

•• Acorales Acorales - sister to all monocots- sister to all monocots•• Alismatids Alismatids

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

–– non-monophyletic non-monophyletic–– petaloid petaloid

•• CommelinidsCommelinids–– Arecales Arecales –– palms palms–– Commelinales Commelinales –– spiderwort spiderwort–– Zingiberales Zingiberales ––bananabanana–– Poales Poales

–– pineapple pineapple–– grasses & sedges grasses & sedges

CommelinidsCommelinids

•• strongly monophyletic! strongly monophyletic!

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

•• this feature allowed placement of this feature allowed placement ofDasypogonaceaeDasypogonaceae

•• largest group of monocots ranging largest group of monocots rangingfrom palms to grassesfrom palms to grasses

CommelinidsCommelinids

DasypogonaceaeDasypogonaceae

4 genera - W Australia4 genera - W Australia

CommelinidsCommelinids•• theme: reduction of flower, loss of theme: reduction of flower, loss ofnectar, loss of zoophily, evolution ofnectar, loss of zoophily, evolution ofbractsbracts

pickeral weedpickeral weed

rapateadrapatead

grassgrass

bromeliadbromeliad

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

•• 190 genera and 2400 species of 190 genera and 2400 species oftrees and shrubstrees and shrubs

•• tropics, subtropics, deserts, tropics, subtropics, deserts,Mediterranean biomesMediterranean biomes

*Arecaceae - palms*Arecaceae - palms•• greatest center of diversity in greatest center of diversity inMalay archipelago, thenMalay archipelago, thenAmazoniaAmazonia

•• depauperate in Africa, but depauperate in Africa, butdiverse in Madagascardiverse in Madagascar

MalaysiaMalaysia

MadagascarMadagascar

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

•• Asian distribution with few species passing through Sulawesi Asian distribution with few species passing through Sulawesior New Guineaor New Guinea

Rattan palm & genericRattan palm & genericdistributionsdistributions

*Arecaceae - palms*Arecaceae - palms

SyagrusSyagrus - lilliput palm of Paraguay - lilliput palm of Paraguay JubaeaJubaea - Chilean wine palm - Chilean wine palm

Great morphological diversity: Great morphological diversity: in staturein stature

*Arecaceae - palms*Arecaceae - palms

Lodoicea maldivicaLodoicea maldivica - Seychelles palm or double nut - Seychelles palm or double nut

Great morphological diversity: Great morphological diversity: largest seed of seed plantslargest seed of seed plants

Lodoicea maldivicaLodoicea maldivica - Seychelles palm or double nut - Seychelles palm or double nut

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

What is unusualWhat is unusualabout how thisabout how thisspecies was firstspecies was firstdiscovered?discovered?

*Arecaceae - palms*Arecaceae - palmsGreat morphological diversity: Great morphological diversity: largest leaflargest leaf

RaffiaRaffia - rattan - rattan. . . and. . . and largest inflorescence largest inflorescence

CoryphaCorypha

RoystoniaRoystonia

*Arecaceae - palms*Arecaceae - palmsVegetative characteristicsVegetative characteristics

•• ““woodywoody”” stems stems via primary via primarythickening meristem or diffusethickening meristem or diffusesecondary growthsecondary growth

•• essentially hardened leaf bases essentially hardened leaf bases

•• single apical meristem: single apical meristem:succeptible to frostsucceptible to frost

*Arecaceae - palms*Arecaceae - palmsVegetative characteristicsVegetative characteristics

•• palmate or pinnate palmate or pinnate compoundcompound,,sheathing, sheathing, plicateplicate or folded or folded

*Arecaceae - palms*Arecaceae - palmsFloral characteristicsFloral characteristics

•• inflorescence surrounded by inflorescence surrounded byspathespathe - once allied with aroids - once allied with aroids


•• flowers unisexual or bisexual flowers unisexual or bisexual

CA 3 CO 3 A 3,6,CA 3 CO 3 A 3,6,∞∞ GG 3 or (3) 3 or (3)


•• fruit a 1-seeded berry or drupe fruit a 1-seeded berry or drupe

ArecaAreca Betel palm Betel palm

CocosCocos coconut coconut

Nypa fruticansNypa fruticanssalt marshsalt marsh

Calamus radicalisCalamus radicalisHawaiiHawaii

*Arecaceae - palms*Arecaceae - palmsClassification: 5 subfamiliesClassification: 5 subfamilies

•• Nypoideae and Calamoideae are first diverging Nypoideae and Calamoideae are first diverging

*Arecaceae - palms*Arecaceae - palmsImportant palms: foodImportant palms: food

Cocos nuciferaCocos nucifera - -coconutcoconut

PhoenixPhoenix - Date palm - Date palm

*Arecaceae - palms*Arecaceae - palmsImportant palms: foodImportant palms: food

ChyrtostachysChyrtostachys - -sealing wax palmsealing wax palm

*Arecaceae - palms*Arecaceae - palmsImportant palms: oil, waxImportant palms: oil, wax

ElaeisElaeis - oil - oilpalmpalm

RoystoneaRoystonea - Royal Palm - Royal Palm WashingtoniaWashingtonia in inSanta MonicaSanta Monica

*Arecaceae - palms*Arecaceae - palmsImportant palms: horticultureImportant palms: horticulture

Palm House at Kew RoyalPalm House at Kew RoyalBotanic Garden - largestBotanic Garden - largestglass house in the worldglass house in the worldwith the largest glass housewith the largest glass houseplant - Chilean wine palmplant - Chilean wine palm

*Arecaceae - palms*Arecaceae - palmsImportant palms: horticultureImportant palms: horticulture

CommelinidsCommelinids4 main groups:4 main groups:






CommelinidsCommelinids•• theme: reduction of flower, loss of theme: reduction of flower, loss ofnectar, loss of zoophily, evolution ofnectar, loss of zoophily, evolution ofbractsbracts


rapateadrapatead

grassgrass

bromeliadbromeliad

Commelinales + ZingiberalesCommelinales + Zingiberales

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


nectarnectar

spiderwortspiderwort

pollen onlypollen only

heliconiaheliconia

nectar + bractsnectar + bracts

Commelinaceae - spiderwortCommelinaceae - spiderwort

Tradescantia ohiensisTradescantia ohiensis - spiderwort - spiderwort

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

Commelina erectaCommelina erecta - Erect dayflower - Erect dayflower



Inflorescence often bractedInflorescence often bracted

RhoeoRhoeo - Moses in a cradle - Moses in a cradle



Flowers actinomorphic orFlowers actinomorphic orzygomorphiczygomorphic

Commelina communisCommelina communis - day flower - day flower

CA 3 CO 3 A 6 CA 3 CO 3 A 6 GG (3) (3)

Commelinaceae - spiderwortCommelinaceae - spiderwort•• species rich in pantropics, species rich in pantropics,especially Africaespecially Africa

•• floral diversity is enormous floral diversity is enormous

Pontederiaceae - pickerel weedPontederiaceae - pickerel weedAquatic family of emergents or floaters.Aquatic family of emergents or floaters.Water hyacinth (Water hyacinth (EichhorniaEichhornia) from) fromtropical America is invasive species intropical America is invasive species insubtropical areas of the world.subtropical areas of the world.

Eichhornia crassipesEichhornia crassipesWater hyacinthWater hyacinthinvading Floridainvading Florida

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

Pontederiaceae - pickerel weedPontederiaceae - pickerel weed

Pontederia cordataPontederia cordata - Pickerel weed - Pickerel weed

Pontederia cordataPontederia cordata - Pickerel weed - Pickerel weed

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

Pontederiaceae - pickerel weedPontederiaceae - pickerel weed Haemodoraceae - kangaroo pawHaemodoraceae - kangaroo paw

AnigozanthusAnigozanthus - kangaroo paw - kangaroo paw

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

ZingiberalesZingiberales•• strongly supported group of 8 tropical strongly supported group of 8 tropicalfamiliesfamilies

•• rhizomatousrhizomatous monocots with showy, monocots with showy,nectared, but highly bracted flowersnectared, but highly bracted flowers

•• 3 shared features: 3 shared features: 1. Parallel-pinnate leaves, often1. Parallel-pinnate leaves, oftendistichousdistichous


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

•• 3 shared features: 3 shared features: 2. Bracted flowers2. Bracted flowersandand inflorescences inflorescences


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

•• 3 shared features: 3 shared features: 3. Inferior ovary3. Inferior ovary

ZingiberalesZingiberales

DNA-based Zingiberales DNA-based Zingiberales ““rhizogramrhizogram”” by John Kress by John Kress

•• order fairly well known based on DNA and morphology order fairly well known based on DNA and morphology

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

CostusCostus floral pattern floral pattern

3 fused sepals3 fused sepals3 separate petals3 separate petals5 fused sterile anthers5 fused sterile anthers(labellum)(labellum)1 fertile anther1 fertile anther

SS SS

SS

PP

PP PPAA

AAAA

AAAAAA

Poales I - showy flowersPoales I - showy flowers4 main groups:4 main groups:






•• showy flowers,showy flowers,insect or birdinsect or birdpollinatedpollinated

•• +/- reduced+/- reducedflowers, insect orflowers, insect orwind pollinatedwind pollinated

•• reducedreducedflowers, windflowers, windpollinatedpollinated

Poales I - showy flowersPoales I - showy flowers

*Bromeliaceae - pineapples*Bromeliaceae - pineapples•• mainly mainly epiphyticepiphytic, but terrestrial as well in, but terrestrial as well ininhospitable regionsinhospitable regions

*Bromeliaceae - pineapples*Bromeliaceae - pineapples•• key adaptations: key adaptations: CAMCAM photosynthesis, modified photosynthesis, modifiedtrichomes or trichomes or scalesscales, , ““tanktank”” formation formation

ScalesScales (water & nutrient(water & nutrientuptake)uptake)

TankTank (water impounding)(water impounding)

*Bromeliaceae - pineapples*Bromeliaceae - pineapples•• key adaptations: key adaptations: CAMCAM photosynthesis, modified photosynthesis, modifiedtrichomes or trichomes or scalesscales, , ““tanktank”” formation formation

•• scales very visible in scales very visible inSpanish mossSpanish moss

Tillandsia usneoidesTillandsia usneoides ininSouth Carolina live oaksSouth Carolina live oaks

*Bromeliaceae - pineapples*Bromeliaceae - pineapples•• preadaptations to preadaptations to carnivorycarnivory in in BrocchiniaBrocchinia and andCatopsisCatopsis

Amino acidsAmino acidsradioactivelyradioactivelylabeled beinglabeled beingincorporated intoincorporated intothe scales ofthe scales ofBrocchiniaBrocchinia

BrocchiniaBrocchinia CatopsisCatopsis

*Bromeliaceae - pineapples*Bromeliaceae - pineapples•• inflorescence heavily bracted and often the inflorescence heavily bracted and often theattractantattractant

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

____

•• petals showy, but not the sepals petals showy, but not the sepals

•• 2 sets of 3 stamens 2 sets of 3 stamens

•• superior or inferior ovary, with superior or inferior ovary, withtwisted stylestwisted styles

•• berry or capsule berry or capsule

*Bromeliaceae - pineapples*Bromeliaceae - pineapples

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

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

guavaguava

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

passion fruitpassion fruit

*Bromeliaceae - pineapples*Bromeliaceae - pineapples•• classification traditionally had three subfamilies classification traditionally had three subfamilies

tillandsioidstillandsioids pitcairnioidspitcairnioids bromelioidsbromelioids

Incan ceremonial danceIncan ceremonial dance

*Bromeliaceae - pineapples*Bromeliaceae - pineapples•• tillandsioids tillandsioids

VrieseaVriesea

Tillandsia usneoides Tillandsia usneoides andand T. grandis T. grandis

*Bromeliaceae - pineapples*Bromeliaceae - pineapples•• bromelioids bromelioids

AchmeaAchmea NeoregeliaNeoregelia

Ananas -Ananas -pineapplepineapple

*Bromeliaceae - pineapples*Bromeliaceae - pineapples•• pitcairnioids pitcairnioids

PuyaPuya

BrocchiniaBrocchinia

NaviaNavia

*Bromeliaceae - pineapples*Bromeliaceae - pineapples•• 2 subfamilies natural 2 subfamilies naturalbased on DNAbased on DNA

•• pitcairnioids broadly pitcairnioids broadlyparaphyleticparaphyletic

•• BrocchiniaBrocchinia sister to rest sister to restof familyof family

•• origin of family in origin of family inGuayana ShieldGuayana Shield of South of SouthAmericaAmerica

*Bromeliaceae - pineapples*Bromeliaceae - pineapplesGuayana HighlandsGuayana Highlands of southern of southernVenezuela and adjacent areas ofVenezuela and adjacent areas ofBrazil and Colombia - the higherBrazil and Colombia - the higherelevation elevation ““tepuistepuis”” are rain are raindrenched and extremely nutrientdrenched and extremely nutrientpoorpoor

*Bromeliaceae - pineapples*Bromeliaceae - pineapplesWhen did theWhen did theAtlantic disjunctionAtlantic disjunctionoccur?occur?

Pitcairnia saxicolaPitcairnia saxicolaCosta RicaCosta Rica

Picairnia felicianaPicairnia feliciana in west Africa in west Africa

BromeliaceaeBromeliaceae

RapateaceaeRapateaceae

DNA tree calibratedDNA tree calibratedwith monocot fossilswith monocot fossils

Long distanceLong distancedispersal to Africa!dispersal to Africa!African speciesAfrican speciesdivergence is 15-13divergence is 15-13myamya

*Bromeliaceae - pineapples*Bromeliaceae - pineapplesWhen did theWhen did theAtlantic disjunctionAtlantic disjunctionoccur?occur?

Atlantic splitAtlantic split

*Bromeliaceae - pineapples*Bromeliaceae - pineapples

•• African speciesAfrican speciesoriginated from Andesoriginated from Andes!!

Where did theWhere did theAfrican speciesAfrican speciescome from?come from?

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

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

•• hummingbird pollination has evolved once in a clade of two hummingbird pollination has evolved once in a clade of twogeneragenera

Is the African Is the African MascolocephalusMascolocephalus a product of Atlantic a product of Atlanticvicariance with closest Guayana Shield relatives, or avicariance with closest Guayana Shield relatives, or a

product of long distance dispersal?product of long distance dispersal?

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

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

Rapateaceae - a tepui familyRapateaceae - a tepui family

Atlantic splitAtlantic split

Commelinids Diversity and Evolution of Monocots · Lodoicea maldivica - Seychelles palm or double...

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