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Diversity and Evolution of Monocots

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Diversity and Evolution of Monocots Commelinids 4 main groups: Diversity and Evolution • Acorales - sister to all monocots • Alismatids of Monocots – inc. Aroids - jack in the pulpit • Lilioids (lilies, orchids, yams) – non-monophyletic . palms, spiderworts, bananas, and – petaloid • Commelinids pineapples . – Arecales – palms – Commelinales – spiderwort – Zingiberales –banana – Poales – pineapple – grasses & sedges Commelinids Commelinids • largest group of monocots ranging from palms to grasses Dasypogonaceae • strongly monophyletic! • bound ferulic acid in cell walls (fluoresce under UV with ammonium hydroxide added) • this feature allowed placement of Dasypogonaceae 4 genera - W Australia Commelinids *Arecaceae - palms • theme: reduction of flower, loss of • the order has one family - also nectar, loss of zoophily, evolution of called Palmae bracts • 190 genera and 2400 species of trees and shrubs • tropics, subtropics, deserts, grass Mediterranean biomes pickeral weed rapatead bromeliad *Arecaceae - palms *Arecaceae - palms Malaysia • greatest center of diversity in • Rattan palms - a plant group that honors the Wallace Malay archipelago, then Biogeographic Line Amazonia • Asian distribution with few species passing through Sulawesi • depauperate in Africa, but or New Guinea diverse in Madagascar Rattan palm & generic distributions Madagascar *Arecaceae - palms *Arecaceae - palms Great morphological diversity: in stature Great morphological diversity: largest seed of seed plants Syagrus - lilliput palm of Paraguay Jubaea - Chilean wine palm Lodoicea maldivica - Seychelles palm or double nut This genus of 1 species endemic to the Seychelles has generated interest in having the largest seed, and in that the shape of the *Arecaceae - palms seed has suggested the devil's work or aphrodisiacal properties. Great morphological diversity: largest leaf What is unusual about how this species was first discovered? Corypha Raffia - rattan Lodoicea maldivica - Seychelles palm or double nut . and largest inflorescence *Arecaceae - palms *Arecaceae - palms Vegetative characteristics Vegetative characteristics • “woody” stems via primary • palmate or pinnate compound, thickening meristem or diffuse sheathing, plicate or folded secondary growth • essentially hardened leaf bases • single apical meristem: succeptible to frost Roystonia *Arecaceae - palms *Arecaceae - palms Floral characteristics Floral characteristics • inflorescence surrounded by • flowers unisexual or bisexual spathe - once allied with aroids CA 3 CO 3 A 3,6,∞∞ G 3 or (3) *Arecaceae - palms *Arecaceae - palms Floral characteristics Areca Betel palm Classification: 5 subfamilies • Nypoideae and Calamoideae are first diverging • fruit a 1-seeded berry or drupe • Nypoideae and Calamoideae are first diverging Cocos coconut Nypa fruticans Calamus radicalis salt marsh Hawaii *Arecaceae - palms *Arecaceae - palms Important palms: food Important palms: food Phoenix - Date palm Cocos nucifera - coconut *Arecaceae - palms *Arecaceae - palms Important palms: oil, wax Important palms: horticulture Elaeis - oil palm Chyrtostachys - Roystonea - Royal Palm Washingtonia in sealing wax palm Santa Monica *Arecaceae - palms Commelinids Important palms: horticulture 4 main groups: Palm House at Kew Royal Botanic Garden - largest • Acorales - sister to all monocots glass house in the world • Alismatids with the largest glass house – inc. Aroids - jack in the pulpit • Lilioids (lilies, orchids, yams) plant - Chilean wine palm • Lilioids (lilies, orchids, yams) plant - Chilean wine palm – non-monophyletic – petaloid • Commelinids – Arecales – palms – Commelinales – spiderwort – Zingiberales –banana – Poales – pineapple – grasses & sedges Commelinids Commelinales + Zingiberales • theme: reduction of flower, loss of nectar, loss of zoophily, evolution of • 2 closely related tropical orders bracts • primarily nectar bearing but with losses • bracted inflorescences grass pickeral weed pickeral weed spiderwort heliconia nectar pollen only nectar + bracts rapatead bromeliad Commelinaceae - spiderwort Commelinaceae - spiderwort Family of small herbs with succulent stems, stems jointed; leaves sheathing. Family does not produce Inflorescence often bracted nectar, but showy flowers for insect pollen gathering. Rhoeo - Moses in a cradle Commelina erecta - Erect dayflower Tradescantia ohiensis - spiderwort Tradescantia ohiensis - spiderwort Commelinaceae - spiderwort Commelinaceae - spiderwort Flowers actinomorphic or • species rich in pantropics, CA 3 CO 3 A 6 G (3) zygomorphic especially Africa • floral diversity is enormous Commelina communis - day flower Tradescantia ohiensis - spiderwort Pontederiaceae - pickerel weed Pontederiaceae - pickerel weed Aquatic family of emergents or floaters. Aquatic family of emergents or floaters. Pickerel weed has glossy heart-shaped leaves, Water hyacinth (Eichhornia) from Water hyacinth (Eichhornia) from superficially like Sagittaria but without net venation. tropical America is invasive species in Flowers are in congested showy purple inflorescences. subtropical areas of the world. Eichhornia crassipes Water hyacinth invading Florida Pontederia cordata - Pickerel weed Pontederiaceae - pickerel weed Haemodoraceae - kangaroo paw Flowers are showy, insect pollinated with nectar glands - previously placed in Liliales! Anigozanthus - kangaroo paw Small family with floral nectar, species radiations in Australia and Pontederia cordata - Pickerel weed South Africa Zingiberales Zingiberales • strongly supported group of 8 tropical • strongly supported group of 8 tropical families families • rhizomatous monocots with showy, • rhizomatous monocots with showy, nectared, but highly bracted flowers nectared, but highly bracted flowers • 3 shared features: 1. Parallel-pinnate leaves, often • 3 shared features: 2. Bracted flowers distichous and inflorescences Zingiberales Zingiberales • strongly supported group of 8 tropical • order fairly well known based on DNA and morphology families • show interesting trends in (1) fusion of perianth and (2) stamen loss and staminode development • rhizomatous monocots with showy, Costus floral pattern nectared, but highly bracted flowers P • 3 shared features: 3. Inferior ovary S A S A A A A P A P S 3 fused sepals 3 separate petals 5 fused sterile anthers (labellum) DNA-based Zingiberales “rhizogram” by John Kress 1 fertile anther Poales I - showy flowers Poales I - showy flowers 4 main groups: • showy flowers, • Acorales - sister to all monocots insect or bird • Alismatids pollinated – inc. Aroids - jack in the pulpit • Lilioids (lilies, orchids, yams) – non-monophyletic • +/- reduced – petaloid flowers, insect or • Commelinids – Arecales – palms wind pollinated – Commelinales – spiderwort – Zingiberales –banana – Poales • reduced – pineapple flowers, wind – grasses & sedges pollinated *Bromeliaceae - pineapples *Bromeliaceae - pineapples • mainly epiphytic, but terrestrial as well in • key adaptations: CAM photosynthesis, modified inhospitable regions trichomes or scales, “tank” formation Tank (water impounding) Scales (water & nutrient uptake) *Bromeliaceae - pineapples *Bromeliaceae - pineapples • key adaptations: CAM photosynthesis, modified • preadaptations to carnivory in Brocchinia and trichomes or scales, “tank” formation Catopsis • scales very visible in Spanish moss Amino acids radioactively labeled being incorporated into Tillandsia usneoides in the scales of South Carolina live oaks Brocchinia Brocchinia Catopsis *Bromeliaceae - pineapples *Bromeliaceae - pineapples __ • inflorescence heavily bracted and often the __ CA 3 CO 3 A 6 G (3) or G (3) attractant • petals showy, but not the sepals • 2 sets of 3 stamens • superior or inferior ovary, with twisted styles • berry or capsule *Bromeliaceae - pineapples *Bromeliaceae - pineapples • bromeliads are an American family: 2600 species, 56 • pineapple not native to Hawaii - along with two other genera ingredients of Hawaiian Punch 1 species of Pitcairnia in west Africa - vicariance or dispersal? guava passion fruit *Bromeliaceae - pineapples *Bromeliaceae - pineapples • classification traditionally had three subfamilies • tillandsioids tillandsioids pitcairnioids bromelioids Tillandsia usneoides and T. grandis Vriesea Incan ceremonial dance *Bromeliaceae - pineapples *Bromeliaceae - pineapples • bromelioids • pitcairnioids Brocchinia Ananas - Navia pineapple Achmea Neoregelia Puya *Bromeliaceae - pineapples *Bromeliaceae - pineapples • 2 subfamilies natural Guayana Highlands of southern based on DNA Venezuela and adjacent areas of Brazil and Colombia - the higher • pitcairnioids broadly elevation “tepuis” are rain paraphyletic drenched and extremely nutrient poor • Brocchinia sister to rest of family • origin of family in Guayana Shield of South America *Bromeliaceae - pineapples *Bromeliaceae - pineapples When did the When did the Atlantic disjunction Atlantic split occur? Atlantic disjunction occur? Bromeliaceae Long distance dispersal to Africa! African species divergence is 15-13 Rapateaceae mya DNA tree calibrated Pitcairnia saxicola Picairnia feliciana in west Africa with monocot fossils Costa Rica *Bromeliaceae - pineapples Rapateaceae - a tepui family • 16 genera and nearly 100 species from the Guayana Shield Where did the African species come from? • African species originated from Andes! Rapateaceae - a tepui family Rapateaceae - a tepui family • most species are pollinated by pollen-gathering bees • most species in the Guayana Shield but one in west Africa • hummingbird pollination has evolved once in a clade of two genera Is the African Mascolocephalus a product of Atlantic vicariance with closest Guayana Shield relatives, or a product of long distance dispersal? Rapateaceae - a tepui family Recent long distance dispersal to Africa! African species divergence is 8-6 my whereas Atlantic separation is 80+ mya Atlantic split.
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