What is native? What is not? When does it matter?

Common milkweed, Asclepias syriaca (© B. Moisset)

Definitions of native plants abound. None is entirely satisfactory in all circumstances but each may serve a specific function. Some strive for scientific accuracy; others serve practical purposes. Moreover, in some instances the nativity of a plant may not matter to the native-plant gardener. None of us is about to give up growing tomatoes regardless of their non-native status.

Some important concepts lack a perfect definition. We need to name them, nonetheless. Biologists would be lost if they couldn’t use the concept of “species” just because no definition fits all the circumstances. This word is not only important but absolutely necessary. One may need to use different definitions of “species” depending on the discipline; but that is no reason to give up the word. The concept of “native” organisms is equally necessary. It is here to stay, regardless of the difficulties that may arise at times.

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© 2012, Beatriz Moisset. All rights reserved. This article is the property of Native Plants and Wildlife Gardens.

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Beginners Guide to Pollinators and Other Flower Visitors

Thrifty flowers

Pollen is a valuable commodity. The less pollen needed, the more energy is left for other functions. Flowers resort to several strategies to economize on pollen. Some use a method called explosive pollination, others resort to buzz pollination. Both methods are well illustrated by official state flowers.

Mountain laurel in bloom. © Beatriz Moisset

Explosive Pollination of Mountain Laurels

Mountain laurel (Kalmia latifolia), Pennsylvania’s and Connecticut's state flower, has a singular way to ensure that its insect visitors carry pollen to other flowers of its species. The unopened blossoms present little knobs which give them a funny look. Their function becomes apparent when the flower opens. They are pockets that hold the anthers trapped. Anthers are the part of the flower that produces male cells, pollen.

Mountain laurel flower before an insect's visit.  © Beatriz Moisset

Flower tripped by an insect. © Beatriz Moisset

In most flowers the anthers are free and exposed to spread their pollen at the slightest touch of a flower visitor. The mountain laurel has a different strategy. The pollen is well protected against rain and wind; but, when a pollinator lands on the flower searching for nectar, the weight acts as a trigger, causing the taut stem of the anther to spring. The anther hits the pollinator gently on the back and gives it a dusting of pollen.

This method allows mountain laurel flowers to produce only a moderate amount of pollen because most of it ends up where it is intended, on the body of a pollinator, rather than being wasted in other ways.

Connecticut: Mountain laurel

Pennsylvania: Mountain laurel

Explosive Pollination of Pea-like Flowers

Texas Bluebonnet. Wikicommons © Jacopo Werther

Other flowers encase both the pollen-producing parts and the female parts, or stigma, into a sheath. Some members of the pea family, Fabaceae, have flowers that resemble a butterfly, this is why this subfamily is called Papilionoideae, papilio meaning butterfly. The top petal is large and shaped as the sail of a ship and is called the banner. Two side petals are called wings, and the bottom two petals are merged into one forming a sheath that encloses both the anthers and the stigma. This structure looks like the keel of a boat and it is so called.

Structure of a papilionoid flower. Wikicommons. © David Richfield

When a pollinator lands on a blossom, its weight triggers a mechanism causing the keel to spread open and the anthers and stigma jump up out of the enclosure. This is called tripping. The insect has to be heavy enough to cause such reaction. Some bumble bees are pros at this task; honey bees, on the other hand, dislike being pounced upon. After a few times, they learn to sneak around and steal the nectar through a small opening at the base of the keel. Only novices perform pollination. Once they learn this trick no more pollen is transferred to the stigmas. Nevertheless managed honey bees are widely used to pollinate alfalfa and clover. Bee hives provide a large labor force that makes up for this deficiency. Some bumble bees, especially the short-tongued species, are also inclined to a little larceny. However, they are generally considered highly competent pollinators of alfalfa and clover.

Clover, another papilionoid flower. © Beatriz Moisset

Texas’ bluebonnet (Lupinus) and Vermont’s red clover (Trifolium pratense) need to be tripped. Six species of lupines, all of them called bluebonnets because of their appearance, live in Texas. All six of them are considered the state flower. The red clover, Vermont’s state flower, on the other hand, is not a native plant. It is an important crop used to feed livestock.

Texas: Bluebonnet

Vermont: Red Clover

Buzz Pollination or the Salt Shaker Technique

Other flowers make the pollinators work for pollen in a different way. Interestingly enough, honey bees never learned how to do the job I will describe, but native bumble bees and numerous species of bees are real pros.

Azalea Andrena bee pollinating azalea. Observe the pores at the tips of the anthers. © Beatriz Moisset

In most flowers the pollen is made available to pollinators as soon as it is ripe. The anther splits open. We all have seen the golden dust many flowers have. If we touch it, it sticks to our fingers. Rhododendrons and azaleas do something else. They keep the pollen enclosed inside the anther. It can only come out through a small opening at the tip. Several other native flowers, like tomato and blueberry have a similar characteristic.

In order to extract it, the bee has to cling to the anther and give it a good shake. It accomplishes this by vibrating its flight muscles while keeping the wings still; it is like running the car engine in neutral. Pollen comes in clouds and clings to the body of the pollinator. Later, the bee proceeds to package its loot in the little baskets of the hind legs.

Buzz pollination of azalea. Video. © Beatriz Moisset

Two states have official flowers that exemplify this method of pollination, Washington has chosen the Coast Rhododendron (Rhododendron macrophyllum) and West Virginia has another species of Rhododendron (Rhododendron maximum). Georgia’s official wildflower, azalea (Rhododendron prunifolium), follows the same system of pollination; its official flower is the rose; but, fortunately, they decided to honor a native plant in addition to a non-native flower.

Georgia (official wildflower): Azalea

Washington: Coast Rhododendron

West Virginia: Rhododendron

Other plants select their pollinator clientele through other methods. Some hide the nectar rather than the pollen. A different set of skills are needed to deal with these flowers as we'll see in the next post.

References

Pollination of mountain laurel

Image illustrating the parts of a Papilionoid flower

Red clover pollination (also alfalfa)

Legume pollination

Bluebonnet pollination

List of articles
Beginners Guide to Pollinators and Other Flower Visitors

© Beatriz Moisset. 2015

Beginners Guide to Pollinators

The Amazon edition of Beginners Guide to Pollinators and Other Flower Visitors came out just in time for Pollinator Week, 2015.

Sample pages below:

This Way to the Restaurant.

Nectar guides © Beatriz Moisset

Nectar Guides

Most flowers want to make it easy for the visitor to find its way around. Thus, in addition to attracting pollinators with their colors and aromas, they guide their visitors in the right direction. This serves two purposes; it facilitates the task of the pollinator and enables the flower to have its pollen deposited where it is most needed.

Have you noticed the streaks of color radiating from the center of many blossoms? These are the nectar guides that tell the visitor: this is the way to the food. Violets and lupines are good examples. The variety of violets is amazing. We can count more than a hundred species and hybrids. Most of them are native, although a few have been introduced from Europe. As a footnote, it is worth mentioning that some violet species are the food plant of fritillary butterflies.

© Beatriz Moisset 

Don’t you find it rather surprising that the humble violet is the state flower of four states? It would be more correct to say violets, plural. Illinois did not attempt to specify which species, even after giving their flower the name of purple violet to distinguish it from yellow violets. New Jersey and Wisconsin chose the Viola sororia and Rhode Island, Viola palmata. Botanists and horticulturists pay attention to these things. However, legislators often ignore such details. This doesn't matter since all violets exemplify the nectar guides vividly.

Illinois: Purple Violet (Viola)

New Jersey: Common Blue Violet (Viola sororia)

Rhode Island: Violet (Viola palmata)

Wisconsin: Common Blue Violet (Viola sororia)

Texas bluebonnet © Jacopo Werther. Wikicommons

A slightly different type of nectar guides is present in bluebonnets (Lupinus). This Texas state flower includes all the species of bluebonnet that grow in that state. The guides become visible only when the flower opens and becomes receptive to pollinators. Bluebonnets belong to the pea family, Fabaceae, which is valuable to ecosystems because it enriches the soil by fixing nitrogen.

Texas: Texas Bluebonnet (Lupinus spp.)

Bull’s Eye

Some flowers use a different way to guide the pollinator to the desired place. It is called bull's eye. The color of the flower's center contrasts with the rest of the blossom. Several state flowers illustrate variations of this theme. Maryland's black-eyed Susan and Oklahoma's Indian blanket are fine examples. So are Florida's tickseed and Kansas' sunflower.

Sunflower © Beatriz Moisset 

Pollination researchers played a trick on visitors to these flowers to test the hypothesis that the bull's eye helped them find their way. They methodically pulled out all the petals of flowers of this type and glued them back in after reversing their position so that the darker part was in the outside. True enough, bees would land on the blossom, walk to the edge and stick their tongues out in search of nectar. They must have been mighty puzzled and annoyed at finding none.

Indian blanket. © Beatriz Moisset 

Some flowers, such as coreopsis, apparently lack this contrast. They appear uniformly yellow to our eyes. But bees see certain colors that escape us. They can see in the range of ultraviolet light, the so-called black light. The flowers, in turn, have a pattern that becomes visible only under this particular kind of light waves. The flower that appears uniformly yellow to us is seen by a bee as having a black center surrounded by a light halo.

Coreopsis. The bull's eye is visible only under ultraviolet light. © Beatriz Moisset 

Maryland, Florida and Kansas have state flowers with bull's eyes, black-eyed Susan, tickseed, and sunflower, respectively. Oklahoma is rather unusual in having not one but three types of flower symbols: the mistletoe as its floral emblem, the Oklahoma rose as its state flower and the Indian blanket, as its state wildflower. The latter has a conspicuous bull's eye.

Maryland: Black-eyed Susan (Rudbeckia hirta)

Florida (Wildflower): Tickseed (Coreopsis)

Kansas: Sunflower (Helianthus annuus)

Oklahoma (Wildflower): Indian Blanket (Gaillardia pulchella)

This article is about one of the many ways in which flowers increase the efficiency of the pollination process. In the next post we'll see examples of another strategy, frugality.

See: Pollinators of Official State Flowers

References

Black-eyedSusan under ultraviolet light

US National Arboretum. Trees and Flower

State Symbols, USA

Netstate. Official State Flowers

Dowden, Anne O. State Flowers. 1978

Cooper, Jason, The Rourke Guide to State Symbols. Flowers. 1942

List of articles
Beginners Guide to Pollinators and Other Flower Visitors

© Beatriz Moisset. 2015

Mass Appeal and Pollination

Some of the many visitors to goldenrod.  © Beatriz Moisset

Many flowers are not particularly selective; they welcome all kinds of visitors. They are wide open, easily accessible, thus many insects can reach the nectar and/or pollen regardless of whether their tongues are long or short. Such flowers provide a convenient standing platform to save their visitors the inconvenience of hovering over them. Their structure is simple so the pollinators don’t need to figure out how to open them in order to reach their rewards. Goldenrod is a fine example.

Monarch butterfly on goldenrod. © Beatriz Moisset

 I have spent countless happy hours observing goldenrods and photographing their numerous visitors. The one most common in my area is the Canada goldenrod. Several hundreds of species visit their flowers. Bear in mind that not all flower visitors qualify as pollinators. Some are not efficient at carrying pollen, or sit in one place for a long time and never make it to another plant. Such is the case of ambush bugs, whose only concern is to snare hapless pollinators to make a meal out of them. If you are interested on photographing and identifying pollinators this is a good place to start.

Metallic green bee. © Beatriz Moisset

 This plant blooms in the fall. It often gets blamed for allergies, but the real culprit is another plant that grows close by, sometimes intermingled with goldenrods, ragweed. Its flowers are inconspicuous because its pollen gets carried by the wind, not by officious pollinators. This is how it gets into our nostrils and causes our suffering. The much maligned goldenrods have larger, heavier and stickier grains of pollen that cannot get airborne and that require help from visitors.

Two states, Kentucky and Nebraska, have the goldenrod as their state flower. It is not clear which species of goldenrod Kentucky picked, but it is likely the Canada goldenrod (Solidago altissima). Nebraska’s choice is clearer, the giant goldenrod (Solidago gigantea). South Carolina, not satisfied with a state flower, also has a state wildflower. It is the Canada goldenrod. It is worth mentioning that Delaware has a state herb, the anise-scented or sweet goldenrod (Solidago odora).

Bumble bee on sunflower. © Beatriz Moisset

Coreopsis. © Beatriz Moisset

 Two other state flowers also have mass appeal. Like goldenrod, they are members of the daisy or aster family, easy to access by different kinds of visitors. They are Kansas’ sunflower (Helianthus annuus) and Maryland’s Black-eyed Susan (Rudbeckia hirta). It is worth mentioning here that two states, Florida and Mississippi, chose the tickseed (Coreopsis spp.) as a state wildflower in addition to their official state flowers, the orange blossom and the magnolia, respectively.

Kentucky: Goldenrod

Nebraska: Goldenrod

South Carolina (wildflower): Goldenrod

Delaware (herb): Goldenrod

Kansas: Sunflower

Maryland: Black-eyed Susan

Florida (wildflower): Coreopsis

Mississippi (wildflower): Coreopsis

The indiscriminate pollination strategy represented by all these flowers has advantages and also disadvantages. They suffer no shortage of pollinators. If one species is doing poorly one year, as it often happens in the insect world, there are plenty more to fill in the deficiency. However, sometimes too much of a good thing can be bad. Some of these non-specialized pollinators may be visiting a variety of flowers instead of being faithful to just one species so they end up carrying the wrong kind of pollen in such cases.

Blossoms with such mass appeal give me the opportunity to introduce the reader to the main groups of pollinators. They are all well represented on these mass appeal flowers.

Andrenidae bee © Beatriz Moisset

Bumble bee. © Beatriz Moisset

 Let us start by getting acquainted with bees. More than 80 species have been seen on goldenrods. Some of them deserve mention. Bumble bees are larger than other bees, plump, dark with yellow or white stripes, and hairy. Metallic green bees are small and gorgeously colored. They may escape notice if one isn’t very observant; but it is easy to become captivated by their shiny aspect. Mason bees and leaf-cutting bees carry pollen on their underbelly, unlike most other bees that carry it on their hind legs. Most of them are dark, nearly black and about the size of honey bees or slightly smaller.

Wasp. © Beatriz Moisset

 Wasps are related to bees and sometimes people confuse them. Most wasps are less hairy than bees and have narrow waists. They raise their young on a diet of insects or spiders, instead of pollen and nectar as bees do. Adult wasps need nectar to fuel their flight. Thus, they are frequent flower visitors and accomplish some pollination.

Syrphid fly. © Beatriz Moisset

Syrphid fly. © Beatriz Moisset

Many types of flies, including mosquitoes and gnats visit flowers to feed on their nectar. Some are good pollinators. The Syrphid flies, also called flower flies, are among the most assiduous flower visitors. Most of them mimic bees or wasps and are frequently mistaken for them. However, they don’t sting and there is no reason to fear them.

Buckeye butterfly on goldenrod. © Beatriz Moisset

Pyrausta moth on sunflower. © Beatriz Moisset

Yellow collared scape moth on goldenrod. © Beatriz Moisset

 Butterflies, skippers and moths have extremely long tongues that they can use like drinking straws when sipping nectar. They favor long necked flowers and carry pollen from one to another.

Polished lady beetle on sunflower. © Beatriz Moisset

Soldier beetle on goldenrod. © Beatriz Moisset

Finally, beetles, although not well specialized for pollination, are also capable of doing so in special circumstances. We already mentioned magnolia pollination by beetles. They also visit other flowers and can do a reasonably good job in some cases.

In the following chapters we’ll see flowers that have chosen a different strategy. They are pickier or more selective. Some attract a handful of visitors; a few go to the extreme of having relationships with only one species of pollinators.

Pollinators of Official State Flowers

References

Goldenrods and Asters: Great PollinatorPlants for the Fall. Penn State Extension

List of articles
Beginners Guide to Pollinators and Other Flower Visitors

© Beatriz Moisset. 2015

The Earliest Pollinators: Beetles and Flies

Magnolia © Beatriz Moisset

 By the time the first flowering plants appeared on Earth there weren’t any bees or butterflies. Those superb pollinators would take millions of years to evolve from wasps and moths respectively. So, who would be attracted to flowers? Who would carry pollen?

Other insects, although not adapted to sipping nectar and storing pollen in little baskets, liked to visit flowering plants to eat the pollen. Sometimes, they also devoured the flowers themselves. Beetles and flies were among the earliest pollinators. These two groups of insects visit the flowers of magnolia and water lilies to this day. In general, flowers pollinated by beetles are cup-shaped to allow these insects to stay for some time. They are strongly scented by fruity or rotten smell. The petals may be tough and leathery, helping them to put up with the abuse; many of them are greenish or creamy white.

Tumbling flower beetles on magnolia © Beatriz Moisset

The state flower of Louisiana and of Mississippi is the Magnolia (Magnolia grandiflora) which is an example of some of the oldest flowering plants; it presents all the qualities listed above and it is pollinated by beetles that use the flowers as a singles bar. They stay for hours eating, drinking, mating and making a mess of the place. When they arrive, usually only the female part of the flower is mature enough, so if they carry pollen from other flowers they get cross-pollinated, but by the time the beetles leave, the stamens or male parts have become ripe. The visitors get easily dusted with it and ready to carry it to the next awaiting singles bar. Beetles and flies find a coating of nectar covering the petals that they can slurp as they go along.

Dance fly on Magnolia © Beatriz Moisset

It is worth mentioning here the tulip tree (Liriodendron tulipifera) because it is the state tree of Indiana, Kentucky and Tennessee. The state flowers of Indiana and Tennessee are not native; they are the peony and iris, respectively, so I like to think of the tulip tree blossom as the honorary state flower of these two states. The tulip tree is a relative of magnolias, equally ancient; its flower bears some resemblance to magnolias. It is also pollinated by beetles, although bees and other insects also contribute to its pollination.

Another ancient flower, the California poppy (Eschscholzia californica), California’s state flower is also pollinated by beetles in some instances. These are more numerous than bees in arid areas. Several species of bees, including honey bees also pollinate these flowers.

California poppy © Audrey. Flickr

State flowers that illustrate the earliest pollinators:
Louisiana: Magnolia

Mississippi: Magnolia

California: California Poppy

Once plants came up with this novel solution to their fertilization process, there was no stopping them. Evolution accelerated and an ever growing variety of flowering plants emerged from the older lineages. In turn, more insects evolved to take advantage of this resource. This is how some carnivorous wasps went vegetarian. They became what we now know as bees. Pollen and nectar supplied all their needs.

Also some moths developed a taste for nectar during their adult life. Unlike most moths, they were diurnal and often sported fancy colors. In other words, they evolved into butterflies. Being frequent flower visitors they became pollinators. This is not to say that wasps and moths, the predecessors of bees and butterflies don’t pollinate. In fact some of them are valuable and are highly specialized ones.

The following posts deal with these pollinators and their flowers.  We will start with the ones who invite a wide assortment of guests. They have mass appeal and several state flowers illustrate this nicely.

Pollinators of Official State Flowers 
Mass Appeal and Pollination

List of articles
Beginners Guide to Pollinators and Other Flower Visitors

© Beatriz Moisset. 2015

Pollinators and vitamins

We need the vitamins provided by fruits

It is said that bees and other pollinators are responsible for one third of all our food. Whether this is exactly right I do not know; but a trip to the grocery store confirms that a substantial part of our food comes from plants that have been pollinated by insects rather than by the wind: most vegetables and fruits, drinks such as coffee and tea. We even have to include in this list beef, poultry and dairy products because farm animals feed partly on alfalfa or clover which have been pollinated by insects. Without pollinators we would be reduced to eating grains or cereals, potatoes, sea food and fish and very little else (and undernourished beef and poultry).

Tomato flowers

What is never mentioned but I find perhaps even more important than food quantity is quality. Many of our essential vitamins and antioxidants come to us courtesy of pollinators. Vegetables and fruits are loaded with vitamins such as beta carotene, vitamin C and a few others.

More evidence can be found in Contribution of Pollinator-Mediated Crops to Nutrients in the Human Food Supply. Elisabeth J. Eilers, Claire Kremen, Sarah Smith Greenleaf, Andrea K. Garber, Alexandra-Maria Klein. The researchers found that most of the vitamins A, C and E come from crops pollinated by insects. A large proportion of the minerals calcium, fluoride and iron in our diet are also dependent on animal pollination. Lycopene and some antioxidants, β-cryptoxanthin and β-tocopherol, are entirely dependent on insect pollination.

So, in summary, if it wasn’t for pollinators we wouldn’t be one third hungrier. Instead we would be one hundred per cent dead.

Vegetables are indispensable because of their vitamins

List of articles
Beginners Guide to Pollinators and Other Flower Visitors

© Beatriz Moisset. 2012

Before there were flowers: Wind Pollination

Pine tassels. © Beatriz Moisset

Pine cones. © Beatriz Moisset

Maine’s state flower is rather unusual because it is not a flower in the real sense. It is the white pine’s (Pinus strobus) tassel and cone. These are not flowers but serve the same function: the tassels produce pollen that has to arrive at the small, immature cones to produce seeds. Only then the cones can grow and reach maturity. Maine residents must be mighty proud of their pines and for very good reasons. It is a handsome tree of great value, and the tassels and cones, although not as colorful as most true flowers, are quite handsome. I am delighted at this peculiar choice because it serves to illustrate a significant point of pollination.

Long before there were any animal pollinators, only the wind performed this function. Wind pollinated plants have to produce vast quantities of pollen so that just a few grains can arrive to their destination; the immense majority never even come near the female cones and simply goes to waste. Pines, firs, spruce, and other conifers belong to an ancient lineage that appeared long before pollinators entered the scene. Thus they were and still are pollinated by the wind. Despite its inefficiency, this method works well enough when many plants of the same species grow relatively close to each other. This is why conifer forests are composed of only a small number of species. This is also why corn, another wind pollinated plant, needs at least several rows of plants to produce seeds.

Pollen dispersal by wind. © Beatriz Moisset

Getting back to the pine, both the tassel and the cone are well-adapted to their functions, increasing the chances that wind-carried pollen finds its destination. The tassels are placed higher on the tree. The cones are aerodynamically shaped to create small air whirlpools that direct the pollen grains toward the seeds. For millions of years, the land was dominated by these plants, along with even older ones, the seedless ones that reproduce by spores. The more ancient dinosaurs never saw a flower, nor did they care. The flower revolution was yet to come, and once started it would take the land by storm and spread to distant corners. Nowadays, wind pollinated conifers are more common than flowering plants only in harsh and cold environments near polar regions. Most plants everywhere else produce flowers.

The great advantage of flowers and of animal pollinated plants is that pollinators are more efficient than wind alone in transporting pollen to the intended target, thus even if individual plants are far apart, they still succeed at being pollinated. In fact, forests of tropical regions are made largely of blooming trees with so many different species packed in the same space that a pollinator has to travel some distance between members of the same species. So pollinators contribute to biodiversity, which in turn contributes to a more efficient way of using all available ecological resources.

Now I can get on with the story of pollinators of state flowers.

Also see: Pollinators of Official State Flowers 

References 
Maine: White pine tassel and cone 

List of articles
Beginners Guide to Pollinators and Other Flower Visitors

© Beatriz Moisset. 2015