Plant Animal

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    Pollination in the Tropics

    1. Pollination BasicsA. Biotic Pollination

    B. Abiotic Pollination

    2. Pollination SyndromesA. Bees

    B. Flies and BeetlesC. Moths

    D. Butterflies

    E. Bats

    F. Birds

    3. Weird Reproduction

    4. Pollination in major

    tropical ecosystems

    5. Humans and Pollination

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    The perfect flower:

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    Pollination basics

    Pollination = the transfer of pollen

    (male microgametophyte) to receptive

    female stigmas for reproduction.

    Plants may be hermaphroditic,

    dioecious or monecious.

    Some plants are self-compatible, while

    others are obligate outcrossers.

    Pollen may be transferred by abiotic

    vectors (wind, water) or biotic vectors

    (animals such as insects, birds, bats).

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    Abiotic Pollination

    No need to attract pollinators

    No production of nectar, no loss of

    pollen, no production of flowers or

    aromas

    The plant can invest all of its energyinto making lots of pollen

    In tropical regions, usually in

    seasonally dry areas, such as in

    savannas where grasses form anherb layer.

    Not very common in rainforests

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    Wind pollination isusually seen in

    grasses, sedges,

    pines, some palm

    trees, and otheropen area species.

    Wind

    pollination

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    Why wind pollination doesnt work well in tropical wet

    forests:

    It is a passive process, and

    depends on environmental

    conditions to be successful

    Lack of consistent windmakes it ineffective in a

    rainforest

    High diversity makes it

    unlikely that a pollen grain will land on a conspecific stigma

    Lack of a leafless stage: its hard for pollen to get around

    High humidity and rainfall reduce amounts of pollen in the air

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    Biotic Pollination

    Mutualistic relationship benefits both the plant and the animal

    Animals: Benefit from an easy source of food (pollen or nectar)

    Get as much as they can at each stop

    Prefer lots of flowers close together, or a few big ones (expend

    less energy on foraging)

    Plants:

    Benefit from dispersal of pollen to as many conspecifics as

    possible: the further the better

    Do not benefit from investing too much in showy flowers,

    scents, attractants

    Benefit from providing as little reward (pollen, nectar) as

    possible for animal consumption

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    250 + MYA

    (late Paleozoic, Mesozoic)

    e.g. Cycadales, seed ferns

    Probably Beetles

    Pollination perhaps inadvertent initially,

    but floral traits that increased chances ofpollination and encouraged visits by most

    efficient pollinators were favored by

    selection

    The carpel (female reproductive structure)

    was originally leaf-shaped. It became foldedon itself to enclose and protect the ovule

    from being eaten by the pollinators. Plants

    with protected ovules would have a

    selective advantage - angiosperms

    Pollination Evolved from Pollinivory in early seed plants

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    2/3 + of Angiosperms are animal pollinated + a smallproportion of Gymnosperms (e.g. Ephedra, cycads)

    Pollinators include:

    Flies (Diptera), Ants, bees, and wasps(Hymenoptera), Beetles (Coleoptera), Butterflies andmoths (Lepidoptera)

    Hummingbirds, sunbirds, honey eaters

    Bats, rodents, small marsupials

    Oddities: Frogs, slugs, earthworms

    Taxonomic Distribution

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    Honey guides

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    Pollination Syndromes

    Often, plants that are visited by particular types ofpollinators will have things in common, especially

    in their flowers, such as certain colors, shapes,

    smells, etc.

    These are generalizations though, not hard and fast rules!

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    PollinatorFlowerColor

    FlowerShape Scent Rewards

    Bees Blue,

    yellow,have UVpattern

    Landing

    platform,corolla tube

    Sweet Usually nectar,

    pollen, resin, orhormoneprecursors

    Beetles Drab Open (flat) Strong; spicy,fruity, or smelly

    Nectar, petals, orspecialized food

    bodiesFlies Drab Open (flat) Strong; spicy or

    smellyMoths Light

    (white or

    cream)

    Tubularcorolla or

    spur

    Strong, sweet;produced in

    evening

    Nectar

    Bats Dull Very longtubes

    Very strongfermenting orfruity

    Lots of nectarand pollen; flowerparts

    Birds Red or

    orange

    Tubular Little or none Nectar, flower

    parts, insects

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    Bees

    Bees are important in all tropical

    communities.In rainforests, bees alone pollinate

    almost half of all canopy trees and

    lianas.

    Bee pollinated flowers:

    are often blue or yellow

    have guide marks that are

    visible in UV light

    have a landing pad

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    Flies and Beetles

    Many fly- or beetle-pollinated plants mimic the

    odor and/or appearance of dung or rotting flesh

    to entice insects for feeding or oviposition.

    They tend to be

    brown or dull-colored.

    Traps and windows are common. There

    is often no nectar. This is called deceit

    pollination.

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    Mostly nocturnal

    Moth pollinated flowers tend to be pale

    Petals tend to be flat or recurved

    Very strong, sweet scent

    May only be open or producing scent or

    nectar at night

    Many moths have extremely long tongues,and feed on nectar from flowers with very

    long corolla tubes, matching the their

    tongue length.

    Some moths that fly in

    the day can be mistaken

    for hummingbirds!

    Moths

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    Butterflies

    Diurnal, good vision - see the color

    red

    Flowers pollinated by diurnal

    butterflies are often colorful andodorless

    Many butterfly flowers occur in

    clusters - Asteraceae, milkweed

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    Bats (Chiropterophily)Nocturnal nectar feeding bats have long

    muzzles and weak teeth, and long tongues

    to scoop pollen. Their flowers are often

    pale, mustily scented, with long tubes and

    wide mouths.

    Pollen from these plants is

    often high in protein and

    important amino acids.

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    Birds

    Bird pollination is extremely important

    in the tropics.Birds are visually oriented, and so their

    flowers often are red, orange, purple or

    yellow.

    They tend to be unscented, with long

    tubes and recurved petals.

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    Pollinator Specialization

    Generalist plants: a variety of smallinsects visit the flowers

    Specialists: the plant is pollinated by a particular

    assemblage of animals. Examples: orchids and

    euglossines,Heliconia and hummingbirds.

    Extreme specialists: almost one-to-one

    host specificity, exemplified by figs and

    fig wasps. This is rare.

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    Freaky Reproduction

    Examples of highly specialized plant-pollinator

    interactions in the tropics are ubiquitous. However, some

    plants have co-evolved with their pollinators to extremes

    that seem almost bizarre.

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    Aroids

    2,500 described species,most pantropical.

    Produce a huge variety of

    scents, from nauseating toblissful.

    Many have heat-

    producing structures: this

    helps diffuse the odors.

    Often pollinated by flies

    or beetles.

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    Some aroids use modified leaves to trap

    pollinators. When insects are attracted to the

    dung-odor ofArum nigrum, they fall into a

    dungeon and are prevented from leaving byone-way filaments. If the insects are carrying

    pollen it is transferred to

    the receptive female

    flowers. Nectar isproduced to feed the

    insects. Twenty-four

    hours later, the male

    flowers mature. The

    filaments shrivel and

    the insects escape -

    but are covered with

    pollen as they exit.

    http://www.youtube.com/watch?v=4P8YhP5_oig&feature=related

    http://www.youtube.com/watch?v=4P8YhP5_oig&feature=relatedhttp://www.youtube.com/watch?v=4P8YhP5_oig&feature=related
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    This Titan Arum can be 12 feet

    tall, and emits a strong smell of

    decay! People used to believe itwas pollinated by elephants.

    The stench attracts insects,

    perhaps carrion beetles, for longdistances to lay their eggs. They

    enter and transfer or receive

    pollen.

    Male and female flowers matureat separate times to avoid self

    pollination.

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    Sauromatum guttatum

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    Over 25,000 species

    described.

    Nearly all orchids are

    pollinated by insects -

    notably bees and wasps,

    but also butterflies, ants,

    flies and others.

    However, almost everyorchid species is

    pollinated by just one or

    two kinds of insects.

    Tight relationship withpollinator: a species of

    orchid may go extinct if

    its pollinator insect dies

    out.

    Orchids

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    Orchids as pollinator traps

    Pollinators ofCoryanthes sp. fit

    exactly to the size of a tunnel

    formed by the lip and the column.

    Approaching male euglossine

    bees are excited by the fragrance,

    and fall into the liquid-filled

    bucket. The only way out is to

    climb up the callus and squeeze

    past the lip, first passing the

    stigma (where pollinia are scrapedoff) and then the pollinarium

    which is stuck onto the body of

    the departing bee.Coryanthes gernotii

    http://www.youtube.com/watch?v=CP1A6TLDQLQ&NR=1

    http://www.youtube.com/watch?v=CP1A6TLDQLQ&NR=1http://www.youtube.com/watch?v=CP1A6TLDQLQ&NR=1
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    Pseudocopulation

    Ophrys is pollinated by several genera ofsolitary bees and wasps. It emits a

    pheromone that mimics the scent of a

    female pollinator, and even looks like one.

    Males are highly attracted to the flowers.

    Their repeated attempts at copulation

    transport the pollinaria between orchids.Often these orchids bloom prior to the

    emergence of the real females.

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    Pseudoaggression

    This orchid (Oncidium planilabre) mimics a male bee.

    When a real bee spots this orchid he attacks it, attemptingto drive the intruder away. He strikes the inflorescence,

    and pollinia are stuck on his head. When he attacks

    another orchid the pollinaria are transferred.

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    Figs and Fig Wasps

    ~1000 species ofFicus (Moraceae), each with own wasppollinator

    fig fruit is a synconium: the inside is full of male and femaleflowers.

    Male and female fig wasp

    (Pleistodontes imperialis). Themale has a greatly reduced body

    with two primary purposes: (1)

    Inseminating the female and (2)

    Drilling exit tunnels through the

    syconium wall.

    V i l i d i i b lli d l d

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    Very precisely timed interaction between pollinators and male andfemale flowers. Female wasps enter a fig with receptive female flowers, ovipositsin sterile flowers and dies.

    Males hatch first, mate with unborn females. Then they chew exittunnels out of the fig before dying. Male flowers are starting tomature and producepollen.

    Newly hatchedfemales leave throughthe tunnels, but pickup pollen in doing so.When they enter a

    new fig they depositthe pollen onreceptive femaleflowers as they searchfor oviposition sites.

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    Pollination in major tropical

    ecosystems Dry Forest - Santa Rosa

    Lowland Rainforest - La Selva

    Moist Forest - BCI

    Montane Forest - Yanayacu

    Mangrove - Milne Bay

    Savannah - Serengeti

    Desert - Sahara

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    Dry forest - Santa Rosa

    Lower species diversity than other

    forest types

    High degree of seasonality

    restricts most pollination activity

    to rainy seasons

    Bees and moths most important

    pollinators, some bats

    Beetle, butterfly, and

    hummingbird pollination are rare

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    Lowland rainforests - La Selva

    Very high diversity of trees -plants are widely distributed, butlocally rare

    Pollination syndromes andcoevolution are common

    Large, showy bracts are common Many tree species are

    outcrossers, some understoryspecies are self-compatible

    Mostly pollinated by bees,especially the canopy

    Flowering time is variable

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    Moist Forest - BCI

    Has a more pronounced

    dry season

    More seasonal flowering

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    Sequential blooming and simultaneous

    fruiting in 6 species ofShorea

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    Montane forests - Yanayacu

    Wet and cold with greaterseasonality than lowland rainforest

    Weather fairly unpredictable

    Lower overall diversity thanlowland rainforest

    Relatively higher diversity ofepiphytes

    Many tree species are self-compatible or dioecious, long

    lasting flowers More hummingbird pollinators,

    fewer bees

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    Mangroves

    Old world mangroves

    tend to be morediverse than those in

    the new world.

    Diverse pollination

    mechanisms,including bats, birds,

    bees and wind.

    Some bear propagules

    rather than seeds;similar to bearing live

    young.

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    Savannah and Desert

    Many plants found in grasslands and deserts

    are wind pollinated, which is possible in

    these open, often windy habitats.

    Vegetation includes perennial

    grasses, shrubs, and trees at

    oases.

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    Desert - Sahara

    Euphorbiaceae - same ecological niche as the

    cacti of the New World deserts produce milky latex

    small flowers in complex inflorescences

    (cacti usually have large single flowers)

    flowers often surrounded by large,

    colorful bracts - pollinator attraction

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    Humans and pollination

    Efficient pollination is critical for agriculture.

    Most food crops require animal pollination Beekeepers have used European honeybees for this in many

    places.

    1/3 of human food is derived from plants pollinated by wildpollinators

    In the US, the value of wild, native pollinators is in thebillions of dollars each year.

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    References:

    Bawa, K.S. 1990. Plant-pollinator interactions in tropical rain

    forests. Annu. Rev. Ecol. Syst. 21:399-422

    Bawa, K.S., and M. Hadley (eds). 1990. Reproductive Ecology of

    Tropical Forest Plants. UNESCO. Kricher, J. 1997. A Neotropical Companion. Princton Univ

    Press.

    Meeuse, B. and S. Morris. 1984. The Sex Life of Flowers.

    Rainbird Pub. Grp. Ltd., London UK

    Proctor, M., P. Yeo, and A. Lack. 1996. The natural history ofpollination. Timber Press, Portland OR.

    Raven, P.H., R.F. Evert, and H. Curtis. 1976. Biology of Plants,

    2nd ed. Worth Pub., New York NY