Diversity and Evolution of Monocots
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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 . spiderworts, bananas, pineapples . – 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 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. Pickerel weed has glossy heart-shaped leaves, 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 Zingiberales Zingiberales Musaceae - banana Musaceae - banana • robust herbs with spiralled phyllotaxy • unisexual flowers • fleshy fruits • “tubular” flowers [3 sepals + 2 petals] • 5 fertile stamens Musa X paradisica (sterile triploid) cultivated banana Zingiberales Zingiberales Strelitziaceae - bird of paradise Strelitziaceae - bird of paradise • woody trunks (usually) with distichous leaves • 3 genera of Gondwanan distribution • 2 fused petals (elaborated flowers for different pollination systems: bird, marsupial, bat) • 5 or 6 fertile stamens Phenakospermum Ravenala Strelitzia Guayana Shield Madagascar South Africa Zingiberales Zingiberales Lowiaceae - orchidantha Heliconiaceae - heliconia • 1 genus • primarily Neotropical • perianth tube = 3 sepals • robust herbs with distichous phyllotaxy • 1 petal = “labellum” • showy bract system • 5 fertile stamens Orchidantha of SE Asia and Pacific Zingiberales Zingiberales Heliconiaceae - heliconia Costaceae - costus • flowers inverted (resupinate) • robust herbs with spiral phyllotaxy • 5 stamens + staminode • double bracted flowers • flower mites Zingiberales Zingiberales Costaceae - costus Zingiberaceae - ginger • two major groups - insect pollinated and bird pollinated • robust herbs with distichous phyllotaxy • “labellum” formed from 5 sterile stamens • ethereal aromatic (ginger) Zingiberales Zingiberales Zingiberaceae - ginger Cannaceae - canna • striking floral diversity and • only Canna of Neotropics pollinators • asymmetrical flowers Zingiberales Poales I - showy flowers Marantaceae - prayer plant 4 main groups: • pantropical, petiolate leaved • Acorales - sister to all monocots • Alismatids • pairs of asymmetrical – inc. Aroids - jack in the pulpit flowers • Lilioids (lilies, orchids, yams) – non-monophyletic – petaloid • Commelinids – Arecales – palms – Commelinales – spiderwort – Zingiberales –banana – Poales – pineapple – grasses & sedges Poales I - showy flowers *Bromeliaceae - pineapples • mainly epiphytic, but terrestrial as well in • showy flowers, inhospitable regions insect or bird pollinated • +/- reduced flowers, insect or wind pollinated • reduced flowers, wind pollinated *Bromeliaceae - pineapples *Bromeliaceae - pineapples • key adaptations: CAM photosynthesis, modified • key adaptations: CAM photosynthesis, modified trichomes or scales, “tank” formation trichomes or scales, “tank” formation • scales very visible in Spanish moss Tank (water impounding) Tillandsia usneoides in South Carolina live oaks Scales (water & nutrient uptake) *Bromeliaceae - pineapples *Bromeliaceae - pineapples • preadaptations to carnivory in Brocchinia and • inflorescence heavily bracted and often the Catopsis attractant Amino acids radioactively labeled being incorporated into the scales of Brocchinia Brocchinia Catopsis *Bromeliaceae - pineapples *Bromeliaceae - pineapples __ • bromeliads are an American family: 2600 species, 56 CA 3 CO 3 A 6 G (3) or G (3) genera • petals showy, but not the sepals • 2 sets of 3 stamens • superior or inferior ovary, with twisted styles • berry or capsule 1 species of Pitcairnia in west Africa - vicariance or dispersal? *Bromeliaceae - pineapples *Bromeliaceae - pineapples • pineapple not native to Hawaii - along with two other • classification traditionally had three subfamilies ingredients of Hawaiian Punch tillandsioids pitcairnioids bromelioids guava passion fruit Incan ceremonial dance *Bromeliaceae - pineapples *Bromeliaceae - pineapples • tillandsioids • bromelioids Ananas - pineapple Tillandsia usneoides and T. grandis Vriesea Achmea Neoregelia *Bromeliaceae - pineapples *Bromeliaceae - pineapples • pitcairnioids • 2 subfamilies natural Brocchinia based on DNA • pitcairnioids broadly paraphyletic • Brocchinia sister to rest of family • origin of family in Navia Guayana Shield of South America Puya *Bromeliaceae - pineapples *Bromeliaceae - pineapples Guayana Highlands of southern When did the Venezuela and adjacent areas of Atlantic disjunction Brazil and Colombia - the higher occur? elevation “tepuis” are rain drenched and extremely nutrient Bromeliaceae poor Rapateaceae Pitcairnia saxicola Picairnia feliciana in west Africa Costa Rica *Bromeliaceae - pineapples *Bromeliaceae - pineapples When did the Where did the Atlantic split Atlantic disjunction African species occur? come from? Long distance dispersal to Africa! • African species African species originated from Andes! divergence is 15-13 mya DNA tree calibrated with monocot fossils Rapateaceae - a tepui family Rapateaceae - a tepui family • 16 genera and nearly 100 species from the Guayana Shield • most species are pollinated by pollen-gathering bees • hummingbird pollination has evolved once in a clade of two genera Rapateaceae - a tepui family Rapateaceae - a tepui family • most species in the Guayana Shield but one in west Africa Recent long distance dispersal to Africa! African species divergence is 8-6 my whereas Atlantic separation is 80+ mya Atlantic split Is the African Mascolocephalus a product of Atlantic vicariance with closest Guayana Shield relatives, or a product of long distance dispersal?.