Introduction COEVOLUTION: PLANTS AND POLLINATORS • > 90% of existing plant species are flowering plants • co-evolution between pollinators and early angiosperms Why so successful? – mutation – natural selection Fossilized flowers 130mya Cross Pollination Cross Pollination Pollination – pollen transfer Outbreeding – fertilization of egg and sperm from genetically different organisms of the same species Bisexual flowers – contain both pollen-producing stamens and ovule-containing carpels Advantages: recessive genes genetic diversity evolutionary flexibility Some plants can self-pollinate Reducing inbreeding: -in some angiosperms, genes expressed in the cells at the stigma surface only allow genetically distinct pollen of the same species to germinate - enzymes inhibit the growth of pollen tube - anthers mature before carpels Question: Why do so many plants cross-pollinate? Anemophily Advantages • Wind pollination Works well in: - open areas - low diversity stands - monocultures - areas of low-moderate rainfall – gymnosperms – approx 10% of angiosperms Maple keys NA hickory tree Mountain cedar 142mya Disadvantages Aquatic Plants - requires windy conditions - requires open spaces - distances are a factor - dry environments better than wet environments - energy costs to producing pollen - approx 30 genera use water to transfer pollen - flowers float at the water’s surface where pollen transfer occurs e.g., Vallisneria ragweed Alnus viridus Animal Pollination Zoophily -A few species are pollinated under water e.g., Ceratophyllum –stamens float to surface where pollen is released - pollen sinks and contacts stigma - many aquatic plants produce flowers above the water - may be wind pollinated or animal pollinated Evolution of Animal Pollination Steps in the evolution of zoophily: - most flowering plants pollinated by animals - mutualism - adaptive radiation of angiosperms - likely evolved from wind pollination and - from exploitative animal-plant interactions ~ 300,000 animal species serve as pollinators - ovules secrete sticky sap to catch wind-carried pollen - sap likely attracted animals - carpel: likely an adaptation to prevent herbivory - plants produced nectar Benefits to Pollinators Benefits to Plants 1. food reward to animal pollinators - pollen - nectar - edible flower petals - consumption of organisms on flowers - no longer dependent on wind - colonization of new environments - can now occupy environments with high species diversity, low wind i.e., rainforests - direct delivery of gametes - enhances outcrossing 2. non-food reward - shelter, shade - compounds for producing phermones Costs to Plants and Pollinators Plants Attract Pollinators Plants - rewards may be costly - attract predators - by flower scent, color, shape, and arrangement - flowers are adapted to attract animals that are best adapted to pollinate them Pollinators - foraging costs - handling costs ~ 700 compounds detected in floral odors Flower odor categories: - Flowery scents - Compounds that mimic sex attractants of insects - Dung or rotten meat smell Plants Attract Pollinators Can Plants Trick Their Pollinators? 1/3 of (30,000) orchids obtain pollination by deception Flower odor categories: Flowery scents: - fragrances result primarily from terpenes, for example: - geraniol (prod by roses) - limonene (citrus plants) - vanillin (vanilla orchids) - eugenol (cloves) Flower color compounds - anthocyanins - anthoxanthins - betalains - carotenoids Sexual deception: mate mimics Conferred by hydrocarbons, alcohols, terpenoids Gorteria diffusa, SA Can pollinators cheat? - visit different species of flowers Male bee-flies try to mate with fuzzy spots on petals Flowers Control Pollinators Pollinator syndromes -By flower shapes and positions - coordinated traits of flowers and pollinators - Landing platforms - deduce pollinator type from the appearance of flowers - used by botanists to develop testable predictions about plant and animal pollination relationships -Nectar guides -Intricate flowers requiring forced entry Pollinator Syndromes Pollinator Syndromes 1. Bees communicate direction and distance of flowers, medium-length tongues so can pollinate a variety of flowers, good color vision, but red-blind, good sense of smell (antennae), “high blossom intelligence” (recognize flowers with large rewards), show constancy (recognize and repeatedly visit favorite flowers) Flowers pollinated by bees: - often bilaterally symmetrical - often blue, purple, lavender, white - landing platform - nectar and pollen 2. Butterflies Diurnal, good color vision, land to feed, “high blossom blossom intelligence”, constancy Flowers pollinated by butterflies: - red, pink, orange, blue colors - bright flowers in clusters - landing platforms Pollinator Syndromes Pollinator Syndromes 3. Moths nocturnal, good sense of smell, may require a landing platform, may hover or land Co-evolution between yucca flowers and yucca moths Flowers pollinated by moths: - often white, light-colored flowers - strongly scented flowers - some species pollinate brightly-colored flowers Night-flying moth with exceptionally long proboscis Pollinator Syndromes 4. Flies: Nectar-feeding flies: - red flowers, little odor, sense sugar through their feet - some flies pollinate sweet-smelling flowers Flowers pollinated by carrion flies: - dull color - strongly scent; carrion, feces - nocturnal Titan - Sumatra Stapelia Rafflesia flower, Borneo Pollinator Syndromes Pollinator Syndromes 5. Beetles 6. Birds: hummingbirds, honeycreepers, lorikeets, sunbirds, parrots - beetle pollination arose prior to the first flowering plants - important pollinators in dry habitats and tropical forests - consumers of ovules, pollen and nectar Flowers pollinated by beetles: - white, purple, yellow, orange - large, flat - easy access - strong fruity odour - thermogenic plants Diurnal, good color vision, poor sense of smell, foraging costs high Flowers pollinated by birds: - open during the day - bright red, deep pink color - place to perch - nectar, pollen Monkey beetles pollinate Sparaxis elegans, SA Carrion beetles African sun bird Pollinator Syndromes Pollinators: Keystone Species • • • 7. Bats species that exert an exceptionally great impact on other species in the ecosystem, relative to its abundance removal of keystone species (pollinators) can have disastrous consequences on ecosystems Bats: important pollinators in the tropics Pteropodid bat Flying Foxes nocturnal, fly far distances, color blind, good sense of smell, require large rewards of nectar and pollen, important pollinators in the tropics Flowers pollinated by bats: - open at night - white-light color - strong fragrances - sturdy flowers - may have landing platform – Widespread on the South Pacific Islands • often the only pollinator and seed disperser – nearly driven to extinction by human hunting, now protected wild banana Pollinator Syndromes Kapok tree: bat pollinated used for fibre Fruits 8. Rodents Flowers pollinated by rodents: - Colchicum: nectar volume and concentration increased at night (Annals of Botany, 2008) - rodents carry pollen on their fur and in feces - seed production dependent on rodent pollinators African lily, Massonia, pollinated by nocturnal rodent, Gerbillurus paeba (Amer. J. Botany, 2001) Colchicum plants, SA - protect seeds from drying out - are attractive and inviting to animals - food source - promote seed distribution - nourish germinating seeds - important nutrition for ½ of world’s population Fruit Dispersers Coevolution is Important in Global Ecology Biodiversity: the existence of flowering plants in nature depends on the presence of appropriate pollinators - Brighamia, Hawaii pollinator extinct