Year of Pollination: Pollination Syndromes and Beyond

A discussion of pollination syndromes should begin with the caveat that they are a largely outdated way to categorize plant-pollinator interactions. Still, they are important to be aware of because they have informed so much of our understanding about pollination biology, and they continue to be an impetus for ongoing research. The concept of pollination syndromes exists in part because we are a pattern seeking species, endeavoring to place things in neat little boxes in order to make sense of them. This is relatively easy to do in a hypothetical or controlled environment where the parameters are selected and closely monitored and efforts are made to eliminate noise. However, the real world is considerably more dynamic than a controlled experiment and does not conform to black and white ways of thinking. Patterns are harder to unveil, and it takes great effort to ensure that observed patterns are genuine and not simply imposed by our pattern seeking brains.

That being said, what are pollination syndromes?  Pollination syndromes are sets of floral traits that are thought to attract specific types of pollinators. The floral traits are considered to have evolved in order to appeal to a particular group of pollinators – or in other words, selective pressures led to adaptations resulting in mutualistic relationships between plants and pollinators. Pollination syndromes are examples of convergent evolution because distantly related plant species have developed similar floral traits, presumably due to similar selection pressures. Pollination syndromes were first described by Italian botanist, Federico Delpino, in the last half of the 19th century. Over several decades his rudimentary ideas were fleshed out by other botanists, resulting in the method of categorization described (albeit briefly) below.

Honey bee on bee's friend (Phacelia tanacetifolia)

A honey bee getting friendly with bee’s friend (Phacelia tanacetifolia)

Pollination by bees (melittophily) – Flowers are blue, purple, yellow, or white and usually have nectar guides. Flowers are open and shallow with a landing platform. Some are non-symmetrical and tubular like pea flowers. Nectar is present, and flowers give off a mild (sometimes strong) sweet scent.

Pollination by butterflies (psychophily) – Flowers are pink, purple, red, blue, yellow, or white and often have nectar guides. They are typically large with a wide landing pad. Nectar is inside a long, narrow tube (or spur), and flowers have a sweet scent.

Pollination by hawkmoths and moths (sphingophily and phalaenophily) – Moth pollinated flowers open at night, have no nectar guides, and emit a strong, sweet scent. Flowers pollinated by hawkmoths are often white, cream, or dull violet and are large and tubular with lots of nectar. Those pollinated by other moths are smaller, not as nectar rich, and are white or pale shades of green, yellow, red, purple, or pink.

Pollination by flies (myophily or sapromyophily) – Flowers are shaped like a basin, saucer, or kettle and are brown, brown-red, purple, green, yellow, white, or blue.  Some have patterns of dots and stripes. If nectar is available, it is easily accessible. Their scent is usually putrid. A sapromyophile is an organism that is attracted to carcasses and dung. Flies that fall into this category visit flowers that are very foul smelling, offer no nectar reward, and essentially trick the fly into performing a pollination service.

Pollination by birds (ornithophily) –  Flowers are usually large, tubular, and red, orange, white, blue, or yellow. They are typically without nectar guides and are odorless since birds don’t respond to scent. Nectar is abundant and found at various depths within the flower.

Pollination by bats (chiropterophily) – Flowers are large, tubular or bell shaped, and white or cream colored with no nectar guides. They open at night, have abundant nectar and pollen, and have scents that vary from musty to fruity to foul.

Pollination by beetles (cantharophily) – Flowers are large and bowl shaped and green or white. There are no nectar guides and usually no nectar. The scent is strong and can be fruity, spicy, or putrid. Like flies, some beetles are sapromyophiles.

Locust borer meets rubber rabbitbrush (Ericameria nauseosa)

A locust borer meets rubber rabbitbrush (Ericameria nauseosa)

In addition to biotic pollination syndromes, there are two abiotic pollination syndromes:

Pollination by wind (anemophily) – Flowers are miniscule and brown or green. They produce abundant pollen but no nectar or odor. The pollen grains are very small, and the stigmas protrude from the flower in order to capture the windborne pollen.

Pollination by water (hydrophily) –  Most aquatic plants are insect-pollinated, but some have tiny flowers that release their pollen into the water, which is picked up by the stigmas of flowers in a similar manner to plants with windborne pollen.

This is, of course, a quick look at the major pollination syndromes. More complete descriptions can be found elsewhere, and they will differ slightly depending on the source. It’s probably obvious just by reading a brief overview that there is some overlap in the floral traits and that, for example, a flower being visited by a bee could also be visited by a butterfly or a bird. Such an observation explains, in part, why this method of categorizing plant-pollinator interactions has fallen out of favor. Studies have been demonstrating that this is not a reliable method of predicting which species of pollinators will pollinate certain flowers. A close observation of floral visitors also reveals insects that visit flowers to obtain nectar, pollen, and other items, but do not assist in pollination. These are called robbers. On the other hand, a plant species may receive some floral visitors that are considerably more effective and reliable pollinators than others. What is a plant to do?

Pollination syndromes imply specialization, however field observations reveal that specialization is quite rare, and that most flowering plants are generalists, employing all available pollinators in assisting them in their reproduction efforts. This is smart, considering that populations of pollinators fluctuate from year to year, so if a plant species is relying on a particular pollinator (or taxonomic group of pollinators) to aid in its reproduction, it may find itself out of luck. Considering that a flower may receive many types of visitors on even a semi-regular basis suggests that the selective pressures on floral traits may not solely include the most efficient pollinators, but could also include all other pollinating visitors and, yes, even robbers. This is an area where much more research is needed, and questions like this are a reason why pollination biology is a vibrant and robust field of research.

A bumble bee hugs Mojave sage (Salvia pachyphylla)

A bumble bee hugs the flower of a blue sage (Salvia pachyphylla)

Interactions between plants and pollinators is something that interests me greatly. Questions regarding specialization and generalization are an important part of these interactions. To help satiate my curiosity, I will be reading through a book put out a few years ago by the University of Chicago Press entitled, Plant-Pollinator Interactions: From Specialization to Generalization, edited by Nickolas M. Waser and Jeff Ollerton. You can expect future posts on this subject as I read through the book. To pique your interest, here is a short excerpt from Waser’s introductory chapter:

Much of pollination biology over the past few centuries logically focused on a single plant or pollinator species and its mutualistic partners, whereas a focus at the level of entire communities was uncommon. Recently we see a revival of community studies, encouraged largely by new tools borrowed from the theory of food webs that allow us to characterize and analyze the resulting patterns. For example, pollination networks show asymmetry – most specialist insects visit generalist plants, and most specialist plants are visited by generalist insects. This is a striking departure from the traditional implication of coevolved specialists!

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