Pollination in the Tropics

1. Pollination Basics
A. B. A. B. C. D. E. F. Biotic Pollination Abiotic Pollination Bees Flies and Beetles Moths Butterflies Bats Birds

2. Pollination Syndromes

3. Weird Reproduction 4. Pollination in major tropical ecosystems 5. Humans and Pollination

The perfect flower:

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).

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 energy into making lots of pollen In tropical regions, usually in seasonally dry areas, such as in savannas where grasses form an herb layer. Not very common in rainforests

Wind pollination
Wind pollination is usually seen in grasses, sedges, pines, some palm trees, and other open area species.

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 wind makes 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

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

Pollination Evolved from Pollinivory in early seed plants

250 + MYA (late Paleozoic, Mesozoic) e.g. Cycadales, seed ferns Probably Beetles Pollination perhaps inadvertent initially, but floral traits that increased chances of pollination and encouraged visits by most efficient pollinators were favored by selection The carpel (female reproductive structure) was originally leaf-shaped. It became folded on itself to enclose and protect the ovule from being eaten by the pollinators. Plants with protected ovules would have a selective advantage - angiosperms

Taxonomic Distribution
2/3 + of Angiosperms are animal pollinated + a small proportion of Gymnosperms (e.g. Ephedra, cycads) Pollinators include: Flies (Diptera), Ants, bees, and wasps (Hymenoptera), Beetles (Coleoptera), Butterflies and moths (Lepidoptera) Hummingbirds, sunbirds, honey eaters Bats, rodents, small marsupials Oddities: Frogs, slugs, earthworms

Honey guides

Pollination Syndromes
Often, plants that are visited by particular types of pollinators 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!

Flower Flower Pollinator Color Shape

Bees Blue, yellow, have UV pattern Drab

Scent

Rewards
Usually nectar, pollen, resin, or hormone precursors Nectar, petals, or specialized food bodies

Landing Sweet platform, corolla tube Open (flat) Strong; spicy, fruity, or smelly Strong; spicy or smelly Strong, sweet; produced in evening Very strong fermenting or fruity Little or none

Beetles

Flies Moths

Drab Light (white or cream) Dull

Open (flat) Tubular corolla or spur Very long tubes Tubular

Nectar

Bats

Birds

Red or orange

Lots of nectar and pollen; flower parts Nectar, flower parts, insects

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

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.

Moths
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!

Butterflies

Diurnal, good vision - see the color red Flowers pollinated by diurnal butterflies are often colorful and odorless Many butterfly flowers occur in clusters - Asteraceae, milkweed

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.

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.

Pollinator Specialization
Generalist plants: a variety of small insects 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.

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.

Aroids
2,500 described species, most pantropical. Produce a huge variety of scents, from nauseating to blissful.

Many have heatproducing structures: this helps diffuse the odors.

Often pollinated by flies or beetles.

Some aroids use modified leaves to trap pollinators. When insects are attracted to the dung-odor of Arum nigrum, they fall into a dungeon and are prevented from leaving by one-way filaments. If the insects are carrying pollen it is transferred to the receptive female flowers. Nectar is produced 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

This Titan Arum can be 12 feet tall, and emits a strong smell of decay! People used to believe it was pollinated by elephants. The stench attracts insects, perhaps carrion beetles, for long distances to lay their eggs. They enter and transfer or receive pollen. Male and female flowers mature at separate times to avoid self pollination.

Sauromatum guttatum

Orchids

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 every orchid species is pollinated by just one or two kinds of insects. Tight relationship with pollinator: a species of orchid may go extinct if its pollinator insect dies out.

Orchids as pollinator traps

Pollinators of Coryanthes 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 scraped off) 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

Pseudocopulation
Ophrys is pollinated by several genera of solitary 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.

Pseudoaggression

This orchid (Oncidium planilabre) mimics a male bee. When a real bee spots this orchid he attacks it, attempting to drive the intruder away. He strikes the inflorescence, and pollinia are stuck on his head. When he attacks another orchid the pollinaria are transferred.

Figs and Fig Wasps

~1000 species of Ficus (Moraceae), each with own wasp pollinator fig fruit is a synconium: the inside is full of male and female flowers.

Male and female fig wasp (Pleistodontes imperialis). The male has a greatly reduced body with two primary purposes: (1) Inseminating the female and (2) Drilling exit tunnels through the syconium wall.

 Very

precisely timed interaction between pollinators and male and female flowers. Female wasps enter a fig with receptive female flowers, oviposits in sterile flowers and dies. Males hatch first, mate with unborn females. Then they chew exit tunnels out of the fig before dying. Male flowers are starting to mature and produce pollen. Newly hatched females leave through the tunnels, but pick up pollen in doing so. When they enter a new fig they deposit the pollen on receptive female flowers as they search for oviposition sites.

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

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

Lowland rainforests - La Selva

Very high diversity of trees plants are widely distributed, but locally rare Pollination syndromes and coevolution are common Large, showy bracts are common Many tree species are outcrossers, some understory species are self-compatible Mostly pollinated by bees, especially the canopy Flowering time is variable

Moist Forest - BCI Has a more pronounced dry season More seasonal flowering

Sequential blooming and simultaneous fruiting in 6 species of Shorea

Montane forests - Yanayacu

Wet and cold with greater seasonality than lowland rainforest Weather fairly unpredictable Lower overall diversity than lowland rainforest Relatively higher diversity of epiphytes Many tree species are selfcompatible or dioecious, long lasting flowers More hummingbird pollinators, fewer bees

Mangroves
Old world mangroves tend to be more diverse 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.

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.

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

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

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 of pollination. Timber Press, Portland OR. Raven, P.H., R.F. Evert, and H. Curtis. 1976. Biology of Plants, 2nd ed. Worth Pub., New York NY