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Year of Pollination: Most Effective Pollinator Principle and Beyond, part two

“The most effective pollinator principle implies that floral characteristics often reflect adaptation to the pollinator that transfers the most pollen, through a combination of high rate of visitation to flowers and effective deposition of pollen during each visit.” – Mayfield, et al., Annals of Botany (2001) 88 (4): 591-596

In part one, I reviewed a chapter by Jose M. Gomez and Regino Zamora in the book Plant-Pollinator Interactions: From Specialization to Generalization that argues that the most effective pollinator principle (MEPP) “represents just one of multiple evolutionary solutions.” In part two, I summarize a chapter by Paul A. Aigner in the same book that further explains how floral characteristics can evolve without strictly adhering to the MEPP.

maximilian sunflower
Aigner is interested in how specialization develops in different environments and whether or not flowering plants, having adapted to interact with a limited number of pollinators, experience trade-offs. A trade-off occurs when a species or population adapts to a specific environmental state and, in the process, loses adaptation to another state. Or in other words, a beneficial change in one trait results in the deterioration of another. Trade-offs and specialization are often seen as going hand in hand, but Aigner argues that trade-offs are not always necessary for an organism to evolve towards specialization. Plant-pollinator interactions provide an excellent opportunity to test this.

“Flowers demand study of specialization and diversification,” Aigner writes, not only due to their ubiquity, “but because much of the remarkable diversity seen in these organisms is thought to have evolved in response to a single and conspicuous element of the environment – pollination by animals.” If pollinators have such a strong influence on shaping the appearance of flowers, pollination studies should be rife with evidence for trade-offs, but they are not. Apart from not being well-studied, Aigner has other ideas about why trade-offs are not often observed in this scenario.

Aigner is particularly interested in specialization occuring in fine-grained environments. A course-grained environment is “one in which an organism experiences a single environmental state for all of its life.” Specialization is well understood in this type of environment. A fine-grained environment is “one in which an organism experiences all environmental states within its lifetime,” such as “a flowering plant [being] visited by a succession of animal pollinators.” For specialization to develop in a fine-grained environment, a flowering plant must “evolve adaptations to a particular type of pollinator while other types of pollinators are also present.”

It’s important to note that the specialization that Aigner mainly refers to is phenotypic specialization. That is, a flower’s phenotype [observable features derived from genes + environment] appears to be adapted for pollination by a specific type of pollinator, but in fact may be pollinated by various types of pollinators. In other words, it is phenotypically specialized but ecologically generalized. Aigner uses a theoretical model to show that specialization can develop in a fine-grained environment with and without trade-offs. He also uses his model to demonstrates that a flower’s phenotype does not necessarily result from its most effective pollinator acting as the most important selection agent. Instead, specialization can evolve in response to a less-effective pollinator “when performance gains from adapting to the less-effective pollinator can be had with little loss in the performance contribution of the more effective pollinator.”

Essentially, Aigner’s argument is that the agents that are the most influential in shaping a particular organism are not necessarily the same agents that offer the greatest contribution to that organism’s overall fitness. This statement flies in the face of the MEPP, and Aigner backs up his argument with (among other examples) his studies involving the genus Dudleya.

Dudleya saxosa (panamint liveforever) - photo credit: wikimedia commons

Dudleya saxosa (panamint liveforever) – photo credit: wikimedia commons

Dudleya is ecologically generalized. Pollinators include hummingbirds, bumblebees, solitary bees, bee flies, hover flies, and butterflies. “Some Dudleya species and populations are visited by all of these taxa, whereas others seem to be visited by only a subset.” Aigner was curious to see if certain species or populations were experiencing trade-offs by adapting to a particular category of pollinators. Aigner found variations in flower characteristics among species and populations as well as differences in pollinator assemblages that visited the various groups of flowers over time but could not conclude that there were trade-offs “in pollination performance.”

In one study, he looked at pollination services provided by hummingbirds vs. bumblebees as corolla flare changed in size. In male flowers, bumblebees were efficient at removing pollen regardless of corolla flare size, while hummingbirds removed pollen more effectively as corolla flare decreased. Both groups deposited pollen more effectively as corolla flare decreased, but hummingbirds more strongly so. Ultimately, Aigner concluded that “the interactions did not take the form of trade-offs,” or, as stated in the abstract of the study, ” phenotypic specialization [for pollination by hummingbirds] might evolve without trading-off the effectiveness of bumblebees.”

Aigner goes on to explain why floral adaptations may occur without obvious trade-offs. One reason is that different groups of pollinators are acting as selective agents for different floral traits, “so that few functional trade-offs exist with respect to individual traits.” Pollinators have different reasons for visiting flowers and flowers use the pollination services of visitors differently. Another reason involves the “genetic architecture” of the traits being selected for. Results can differ depending on whether or not the genes being influenced are linked to other genes, and genetically based fitness trade-offs may not be observable phenotypically. Further studies involving the genetic architecure of specialized phenotypes are necessary.

And finally, as indicated in part one, pollinators are not the only floral visitors. In the words of Aigner, “if floral larcenists and herbivores select for floral traits in different directions than do pollinators, plants may face direct trade-offs in improving pollination service versus defending against enemies.” These “floral enemies” can have an effect on the visitation rates and per-visit effectiveness of pollinators, which can drastically alter their influence as selective agents.

Like pollination syndromes, the most effective pollinator principle seems to have encouraged and directed a huge amount of research in the field of pollination biology, despite not holding entirely true in the real world. As research continues, a more complete picture will develop. It doesn’t appear that it will conform to an easily digestible principle, but there is no question that, even in its complexity, it will be fascinating.

I will end as I began, with an excerpt from Thor Hanson’s book, The Triumph of Seeds: “The notion of coevolution implies that change in one organism can lead to change in another – if antelope run faster, then cheetahs must run faster still to catch them. Traditional definitions describe the process as a tango between familiar partners, where each step is met by an equal and elegant counter-step. In reality, the dance floor of evolution is usually a lot more crowded. Relationships like those between rodents and seeds [or pollinators and flowers] develop in the midst of something more like a square dance, with couples constantly switching partners in a whir of spins, promenades, and do-si-dos. The end result may appear like quid pro quo, but chances are a lot of other dancers influenced the outcome – leading, following, and stepping on toes along the way.”

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Year of Pollination: Most Effective Pollinator Principle and Beyond, part one

Have you ever considered the diversity of flowers? Why do they come in so many different shapes, sizes, and colors? And why do they produce so many different odors – or none at all? Flowering plants evolved around 140 million years ago, a fairly recent emergence evolutionarily speaking. Along with them evolved numerous species of insects, birds, and mammals. In his book, The Triumph of Seeds, Thor Hanson describes the event this way: “In nature, the flowering plants put sex, seeds, and dispersal on full display, spurring not only their own evolution but also that of the animals and insects with which they became so entwined. In most cases, the diversity of dispersers, consumers, parasites – and, most especially, pollinators – rose right alongside that of the plants they depended upon.”

Speaking of dependence, most flowering plants depend upon pollinators for successful reproduction – it is, for the most part, a mutually beneficial relationship. Even the casual observer of flowers will note that a large portion of the creatures that visit them appear to be pollinators. Thus, it is no wonder that pollination biologists have given pollinators so much credit in shaping the flowers that we see today.

Consider G. Ledyard Stebbins and his Most Effective Pollinator Principle which he defined in a paper published in 1970: “the characteristics of the flower will be molded by those pollinators that visit it most frequently and effectively in the region where it is evolving.” He then goes on to reference pollination syndromes, a phenomenon that describes how the traits of flowers are best suited for their “predominant and most effective vector[s].” In my post about pollination syndromes a few months ago, I discussed how a strict adherence to this concept has waned. In the next two posts, I discuss how the Most Effective Pollinator Principle (MEPP) may not be the best way to explain why flowers look the way they do.

 

To make this argument I am drawing mainly from two chapters in the book Plant-Pollinator Interactions: From Specialization to Generalization. The first is “Ecological Factors That Promote the Evolution of Generalization in Pollination Systems” by Jose M. Gomez and Regino Zamora, and the second is “The Evolution of Specialized Floral Phenotypes in a Fine-grained Pollination Environment” by Paul A. Aigner.

According to Aigner the MEPP “states that a plant should evolve specializations to its most effective pollinators at the expense of less effective ones.” And according to Gomez and Zamora it “states that natural selection should modify plant phenotypes [observable characteristics derived from interactions between a plant’s genes and its surrounding environment] to increase the frequency of interaction [between] plants and the pollinators that confer the best services,” and so “we would expect the flowers of most plants to be visited predominantly by a reduced group of highly effective pollinators.” This is otherwise known as adaptive specialization.

Specialization is something that, in theory, plants are generally expected to evolve towards, particularly in regards to plant-pollinator relationships. Observations, on the other hand, demonstrate the opposite – that specialization is rare and most flowering plants are generalists. However, the authors of both chapters advise that specialization and generalization are extreme ends to a continuum, and that they are comparative terms. One species may be more specialized than another simply because it is visited by a smaller “assemblage” of pollinators. The diversity of pollinators in that assemblage and the pollinator availability in the environment should also be taken into consideration when deciding whether a relationship is specialized or generalized.

That pollinators can be agents in shaping floral forms and that flowering plant species can become specialized in their interactions with pollinators is not the question. There is evidence enough to say that it occurs. However, that the most abundant and/or effective pollinators are the main agents of selection and that specialization is a sort of climax state in the evolutionary process (as the MEPP seems to suggest) is up for debate. Generalization is more common than specialization, despite observations demonstrating that pollinators are drawn to certain floral phenotypes. So, could generalization be seen as an adaptive strategy?

In exploring this question, Gomez and Zamora first consider what it takes for pollinators to act as selective agents. They determine that “pollinators must first benefit plant fitness,” and that when calculating this benefit, the entire life cycle of the plant should be considered, including seed germination rate, seedling survival, fecundity, etc. The ability of a pollinator species to benefit plant fitness depends on its visitation rate and its per-visit effectiveness (how efficiently pollen is transferred) – put simply, a pollinator’s quantity and quality during pollination. There should also be “among-pollinator differences in the evolutionary effect on the plant,” meaning that one species or group of pollinators – through being more abundant, effective, or both – contributes more to plant fitness compared to others. “Natural selection will favor those plant traits that attract the most efficient or abundant pollinators and will also favor the evolution of the phenotypes that cause the most abundant pollinators to also be the most effective.” This process implies possible “trade-offs,” which will be discussed in part two.

When pollinators act as selective agents in this way, the MEPP is supported; however, Gomez and Zamora argue that this scenario “only takes place when some restrictive ecological conditions are met” and that while specialization can be seen as the “outcome of strong pollinator-mediated selection,” generalization can also be “mediated by selection exerted by pollinators…in some ecological scenarios.” This is termed adaptive generalization. In situations where ecological forces constrain the development of specialization and pollinators are not seen as active selection agents, nonadaptive generalization may be occurring.

Gomez and Zamora spend much of their chapter exploring “several causes that would fuel the evolution of generalization” both adaptive and nonadaptive, which are outlined briefly below.

  • Spatiotemporal Variability: Temporal variability describes differences in pollinator assemblages over time, both throughout a single year and over several years. Spatial variability describes differences in pollinator assemblages both among populations where gene flow occurs and within populations. Taken together, such variability can have a measurable effect on the ability of a particular pollinator or group of pollinators to act as a selective agent.
  • Similarity among Pollinators: Different pollinator species can have “equivalent abundance and above all comparable effectiveness” making them “functional equivalents from the plant perspective.” This may be the case with both closely and distantly related species. Additionally, a highly effective pollinator can select for floral traits that attract less effective pollinators.
  • The Real Effects on Plant Fitness: An abundant and efficient pollinator may select for one “fitness component” of a plant, but may “lead to a low overall effect on total fitness.” An example being that “a pollinator may benefit seed production by fertilizing many ovules but reduce seedling survival because it causes the ripening of many low-quality seeds.” This is why “as much of the life cycle as possible” should be considered “in assessing pollinator effectiveness.”
  • Other Flower Visitors: Pollinators are not the only visitors of flowers. Herbivores, nectar robbers, seed predators, etc. may be drawn in by the same floral traits as pollinators, and pollinators may be less attracted to flowers that have been visited by such creatures. “Several plant traits are currently thought to be the evolutionary result of conflicting selection exerted by these two kinds of organisms,” and “adaptations to avoid herbivory can constrain the evolution of plant-pollinator interactions.”

This, of course, only scratches the surface of the argument laid out by Gomez and Zamora. If this sort of thing interests you, I highly encourage you to read their chapter. Next week I will summarize Aigner’s chapter. If you have thoughts on this subject or arguments to make please don’t hesitate to comment or contact me directly. This is a dialogue, dudes.

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Year of Pollination: Pollinator Walk at Earthly Delights Farm

Last week I had the privilege of attending a pollinator walk with a local entomologist at Earthly Delights Farm, a small, urban farm in Boise, Idaho. The entomologist was Dr. Karen Strickler, an adjunct instructor at College of Western Idaho and the owner of Pollinator Paradise. A small group of us spent a couple of hours wandering through the farm looking for pollinators and discussing whatever pollinator or non-pollinator related topic that arose. Earthly Delights Farm, along with growing and selling produce using a subscription-based model, is a seed producing farm (and part of a larger seed growing operation called Snake River Seed Cooperative), so there were several crops flowering on the farm that would typically be removed at other farms before reaching that stage, such as lettuce and carrots. The farm also shares property with Draggin’ Wing High Desert Nursery, a nursery specializing in water efficient plants for the Intermountain West, which has a large demonstration area full of flowering plants. Thus, pollinators were present in abundance.

A series of isolation tents over various crops to help prevent cross pollination between varieties.

A series of isolation tents placed over various crops to help prevent cross pollination between varieties – an important component of seed saving.

While many groups of pollinators were discussed, including leafcutter bees, bumblebees, honeybees, sweat bees, hummingbirds, and beetles, much of our conversation and search was focused on syrphid flies. Flies are an often underappreciated and overlooked group of pollinators. While not all of the 120,000 species of flies in the world are pollinators, many of them are. The book Attracting Native Pollinators by the Xerces Society has this to say about flies: “With their reputation as generalist foragers, no nests to provision, and sometimes sparsely haired bodies, flies don’t get much credit as significant pollinators. Despite this reputation, they are often important pollinators in natural ecosystems for specific plants, and occasionally for human food plants.” They are especially important pollinators in the Arctic and in alpine regions, because unlike bees, they do not maintain nests, which means they use less energy and require less nectar, making them more fit for colder climates.

One food crop that flies are particularly efficient at pollinating is carrots. According the Xerces Society, carrot flowers are “not a favorite of managed honeybees.” Most flies do not have long tubular, sucking mouthparts, so they search for nectar in small, shallow flowers that appear in clusters, such as plants in the mint, carrot, and brassica families. Flower-visiting flies come in search of nectar and sometimes pollen for energy and reproduction. While acquiring these meals they can at times inadvertently collect pollen on their bodies and transfer it to adjacent flowers. They are generally not as efficient at moving pollen as other pollinators are, but they can get the job done.

Blister beetle on carrot flowers (a preferred food source of flies). Beetles can be important pollinators, even despite chewing on the flowers as they proceed.

Blister beetle on carrot flowers (a preferred food source of flies). Beetles can be effective pollinators as well, even despite chewing on the flowers as they proceed.

During the pollinator walk, we were specifically observing flies in the family Syrphidae, which are commonly known as flower flies, hoverflies, or syrphid flies. Many flies in this family mimic the coloring of bees and wasps, and thus are easily confused as such. Appearing as a bee or wasp is a form of protection from predators, who typically steer clear from these insects to avoid being stung. The larvae of syrphid flies often feed on insects, a trait that can be an added benefit for farmers and gardeners, particularly when their prey includes pest insects like aphids. Other families of flies that are important pollinators include Bombyliidae (bee flies), Acroceridae (small-headed flies), Muscidae (house flies), and Tachinidae (tachinid flies).

Common banded hoverfly (Syrphus ribesii) - one species of hundreds in the syrphid fly family, a common and diverse family of flower visiting flies (photo credit: www.eol.org)

Common banded hoverfly (Syrphus ribesii) – one species of thousands in the syrphid fly family, a common and diverse family of flower-visiting flies (photo credit: www.eol.org)

Because many species of flies visit flowers and because those flies commonly mimic the appearance of bees and wasps, it can be difficult to tell these insects apart. Observing the following features will help you determine what you are looking at.

  • Wings – flies have two; bees have four (look closely though because the forewings and hindwings of bees are attached with a series of hooks called hamuli making them appear as one)
  • Hairs – flies are generally less hairy than bees
  • Eyes – the eyes of flies are usually quite large and in the front of their heads; the eyes of bees are more towards the sides of their heads
  • Antennae – flies have shorter, stubbier antennae compared to bees; the antennae of flies also have bristles at the tips
  • Bees, unlike flies, have features on their legs and abdomens designed for collecting pollen; however, some flies have mimics of these features
Bumblebee on Echinacea sp.

Bumblebee visiting Echinacea sp.

Another interesting topic that Dr. Strickler addressed was the growing popularity of insect hotels – structures big and small that are fashioned out of a variety of natural materials and intended to house a variety of insects including pollinators. There is a concern that many insect hotels, while functioning nicely as a piece of garden artwork, often offer little in the way of habitat for beneficial insects and instead house pest insects such as earwigs. Also, insect hotels that are inhabited by bees and other pollinators may actually become breeding grounds for pests and diseases that harm these insects. It is advised that these houses be cleaned or replaced regularly to avoid the build up of such issues. Learn more about the proper construction and maintenance of insect hotels in this article from Pacific Horticulture.

A row of onions setting seed at Earthly Delights Farm. Onions are another crop that is commonly pollinated by flies.

A row of onions setting seed at Earthly Delights Farm. Onions are another crop that is commonly pollinated by flies.

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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!

References:

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How Pitcher Plants Eat Bugs (Frog Optional)

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A few months ago at work I captured this photo of a frog inside of a pitcher plant. Do you see it? It is pretty well camouflaged and poking its head out just enough to intercept curious insects lured in by the promise of nectar, eating them before they can make their way into the tube. Either way, approaching insects are about to meet their fate. Whether by plant or by frog, they are destined to be consumed lest they turn away in time.

This frog was hiding inside the modified leaf of a species of Sarracenia, a carnivorous plant commonly known as a North American pitcher plant. There are at least eight species of Sarracenia, all of which naturally occur in the southeastern region of the United States. One species, Sarracenia purpurea, also occurs in the northeast, the upper Midwest, and throughout much of Canada. Sarracenia is in the family Sarraceniaceae along with two other genera of pitcher plants, Darlingtonia (the cobra plant, native to northern California and southern Oregon) and Heliamphora (the sun pitchers, native to South America). Plants in this family are not to be confused with the distantly related tropical pitcher plants which are in the genus Nepenthes (family Nepentheaceae).

The natural habitats of Sarracenia are sunny, open areas that remain permanently wet, including meadows, savannahs, fens, and swamps. The soils are acidic, nutrient poor, and typically composed of sandy peat commonly derived from sphagnum moss. In the southeast, less than 5% of the original (pre-European settlement) Sarracenia habitat remains, threatening its survival in the wild. Sarracenia oreophila (green pitcher plant) is currently listed as critically endangered on the IUCN Red List.

Flowering occurs in the spring, usually before pitchers form. Individual flowers are formed on tall stalks that rise straight up and then bend at the very top, hanging the flower upside down. Early flowering and tall flower stalks help prevent pollinating insects from being consumed by the plant. In his book The Savage Garden, Peter D’Amato describes the flowers as “showy, brilliant, and very unusual – a wonderful bonus to an already handsome class of foliage plants.” The flowers are either yellow or a shade of red and last about two weeks, after which the petals drop and a seed pod forms. Seeds are released from the fruits in the fall.

Flower of Sarracenia rubra (sweet pitcherplant) - photo credit: www.eol.org

Flower of Sarracenia rubra (sweet pitcher plant) – photo credit: www.eol.org

D’Amato writes that Sarracenia are among the “most ravenous” plants, with each leaf having the potential of trapping “thousands of nasty insects.” In some cases pitchers even flop over, heavy with the weight of bugs inside them. The specifics of capturing and killing insects varies between species of Sarracenia, but in general prey is lured to the opening of the pitcher with a combination of nectar, scent, and color. Upon entering the tube, gravity, waxy surfaces, drugs, and hairs force the captives downward where they are eventually consumed by enzymes and microbes. Digested insects provide the plant with nutrients necessary for growth – nutrients that otherwise are taken up by the roots of plants that occur in more nutrient rich soils.

Sarracenia purpurea (purple pitcher plant) is unique in that its pitchers lack a “hood” or “lid” – a standard feature of other species of Sarracenia that helps keep rain from entering the pitchers. Instead, the pitchers fill with water and insects are killed by drowning. The most brutal killer is probably Sarracenia psittacina (parrot pitcher plant) which has an additional opening inside of its pitcher. The opening is small and difficult to find again once an insect is on the wrong side of it. The inside walls of the pitcher are covered in long, sharp, downward pointing hairs, and the struggling insect is pierced repeatedly by the hairs as it makes its way to the bottom of the tube to be digested.

Hoodless pitchers of Sarracenia purpurea (photo credit: www.eol.org)

Hoodless pitchers of Sarracenia purpurea (photo credit: www.eol.org)

Hooded pitchers of Sarracenia leucophylla (photo credit: www.eol.org)

Hooded pitchers of Sarracenia leucophylla (photo credit: www.eol.org)

According to D’Amato, “the Sarracenia are one of the simplest carnivorous plants to grow, and certainly among the most fun and rewarding.” Learn more about growing North American pitcher plants by consulting D’Amato’s book and/or by visiting the website of the International Carnivorous Plant Society.

Want to learn more about Sarracenia? The Plants are Cool, Too! web series has a great video about them:

Other carnivorous plant posts:

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Year of Pollination: The Anatomy of a Bee

A greater appreciation for pollinators can be had by learning to identify them – being able to tell one from another and calling them by name. Anyone can tell a butterfly from a bee, but how about telling a sweat bee from a leafcutter bee? Or one species of sweat bee from another species of sweat bee? That takes more training. This is where knowing the parts of a bee becomes important.

I am new to learning the names of pollinators. I’ve been learning the names of plants for many years now (and I still have a long way to go), but my knowledge of insect identification is largely limited to one entomology course I took in college and the occasional reading about insects in books and magazines. So, this post is just as much for me as it is for anybody else. It also explains why it is brief and basic. It’s for beginners.

This first illustration is found in the book Pollinators of Native Plants by Heather Holm. The book starts with brief overviews of pollination, pollinators, and pollinator conservation, but then spends nearly 200 pages profiling specific plants and describing the particular species of pollinating insects that visit them. The photos of the insects are great and should be very useful in helping to identify pollinators.

bee anatomy_pollinators of native plants book

This next illustration is from the book California Bees and Blooms by Gordon W. Frankie, et al. The title is a bit deceptive because much of what is found in this book is just as applicable to people outside of California as it is to people within. There is some discussion about plants and pollinators specific to California and the western states, but there is also a lot of great information about bees, flowers, and pollination in general, including some great advice on learning to identify bees. The book includes this basic diagram, but it also provides several other more detailed illustrations that help further describe things like mouth parts, wings, and legs.

bee anatomy_california bees and blooms book

As part of their discussion on identifying bees, the authors of California Bees and Blooms offer these encouraging and helpful words to beginners like me: “Even trained taxonomists must examine most bees under a microscope to identify them to species level, but knowing the characteristics to look for can give you a pretty good idea of the major groups and families of bees that are visiting your garden. These include size, color, and features of the head, thorax, wings, and abdomen.”

If you would like to know more about the pollinators found in your region, including their names, life history, and the plants they visit, books like the aforementioned are a good start. Also, find yourself a copy of a field guide for the insects in your area and a good hand lens. Then spend some time outside closely and quietly observing the busy lives of the tiny things around you. I plan to do more of this sort of thing, and I am excited see what I might find. Let me know what you find.

Here are a few online resources for learning more about bee anatomy and bee identification:

Other “Year of Pollination” Posts:

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Plants Use Mycorrhizal Fungi to Warn Each Other of Incoming Threats

The March 2015 issue of New Phytologist is a Special Issue focusing on the “ecology and evolution of mycorrhizas.” This is the second of two articles from that issue that I am reviewing. Read the first review here.

Interplant signalling through hyphal networks by David Johnson and Lucy Gilbert

When an individual plant is attacked by an insect or fungal pest, it can warn neighboring plants – prompting them to produce compounds that either repel the pests or attract beneficial organisms that can fight off the pests. There are two main pathways for a plant to send these communications: through the air by way of volatile organic compounds (VOC’s) or through the soil by way of a vast collection of fungal hyphae called mycelium. Plant communication by aerial release of VOC’s has been well documented; communication via mycelium, however, is a fairly recent discovery, and there is much left to learn.

“The length of hyphae in the soil and the ability of mycorrhizal fungi to form multiple points of entry into roots can lead to the formation of a common mycelial network (CMN) that interconnects two or more plants.” These CMN’s are known to play “key roles in facilitating nutrient transport and redistribution.” We now understand that they can also “facilitate defense against insect herbivores and foliar necrotophic fungi by acting as conduits for interplant signaling.” The purpose of this research is to explore the “mechanisms, evolutionary consequences, and circumstances” surrounding the evolution of this process and to “highlight key gaps in our understanding.”

interplant signaling

An illustration of plant communication (aka interplant signaling) by air and by soil form the article in New Phytologist.

If plants are communicating via CMN’s, how are they doing it? The authors propose three potential mechanisms. The first is by signal molecules being transported “in liquid films on the external surface of hyphae via capillary action or microbes.” They determine that this form of communication would be easily disrupted by soil particles and isn’t likely to occur over long distances. The second mechanism is by molecules being transported within hyphae, passing from cell to cell until they reach their destination. The third mechanism involves an electrical signal induced by wounding.

If signal molecules are involved in the process, what molecules are they? There are some molecules already known to be transported by fungal hyphae (lipids, phosphate transporters, and amino acids) and there are also compounds known to be involved in signaling between plants and mycorrhizal fungi. Exploring these further would be a good place to start. We also need to determine if specific insect and fungal pests or certain types of plant damage result in unique signaling compounds.

It has been established that electrical signals can be produced in response to plant damage. These signals are a result of a process known as membrane depolarization. “A key advantage of electrical-induced defense over mobile chemical is the speed of delivery.” Movement of a molecule through cells occurs significantly slower than an electrical charge, which is important if the distance to transport the message is relatively far and the plant needs to respond quickly to an invasion. Various aspects of fungal physiology and activity have been shown to be driven in part by membrane depolarization, so involving it in interplant signaling isn’t too far-fetched.

photo credit: wikimedia commons

photo credit: wikimedia commons

How and why does a system of interplant communication involving mycorrhizal fungi evolve? And what are the costs and benefits to the plants and fungi involved? Determining costs and benefits will depend largely on further establishing the signaling mechanisms. Exploring real world systems will also help us answer these questions. For example, in a stable environment such as a managed grassland where CMNs are well developed, a significant loss of plants to a pest or disease could be devastating for the mycorrhizal community, so “transferring warning signals” would be highly beneficial. Conversely, in an unstable environment where a CMN is less established, assisting in interplant signaling may be less of an imperative. Regarding questions concerning the degree of specialization involved in herbivore-plant-fungal interactions: if a “generic herbivore signal” is sent to a neighboring plant that is not typically affected by the attacking herbivore, the cost to the plant in putting up its defenses and to the fungus in transporting the message is high and unnecessary. So, in an environment where there are many different plant species, species-specific signals may be selected for over time; in areas where there are few plant species, a generic signal would suffice.

As research continues, the mysteries of “defense-related” interplant communication via CMN’s will be revealed. Field studies are particularly important because they can paint a more accurate picture compared to “highly simplified laboratory conditions.” One exciting thing about this type of communication is that it may mean that some plants are communicating with each other across great distances, since “some species of fungi can be vast.” CMNs can also target specific plants, and compared to communication via aerial VOC’s, the signal will not be diluted by the wind.

Since I am in the process of reading Robin Wall Kimmerer’s book, Braiding Sweetgrass, I have decided to include her description of a tree-mycorrhizal fungi relationship:

The trees in a forest are often interconnected by subterranean networks of mycorrhizae, fungal strands that inhabit tree roots. The mycorrhizal symbiosis enables the fungi to forage for mineral nutrients in the soil and deliver them to the tree in exchange for carbohydrates. The mycorrhizae may form fungal bridges between individual trees, so that all the trees in a forest are connected. These fungal networks appear to redistribute the wealth of carbohydrates from tree to tree. A kind of Robin Hood, they take from the rich and give to the poor so that all the trees arrive at the same carbon surplus at the same time. They weave a web of reciprocity, of giving and taking. In this way, the trees all act as one because the fungi have connected them. Through unity, survival. All flourishing is mutual.

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Year of Pollination: Dung Moss

Last year I wrote about two groups of plants that emit foul odors when they bloom: corpse flowers and carrion flowers. Their scent is akin to the smell of rotting flesh, hence their common names. The purpose of this repugnant act is to attract a specific group of pollinators: flies, carrion beetles, and other insects that are attracted to gross things. Though this particular strategy is rare, these aren’t the only plants that have evolved to produce stinky smells in order to recruit such insects to aid in their reproductive processes. For one, there is a very unique group of mosses that do this, commonly known as dung mosses. Judging from the name, you can probably imagine what their smell must be like. However, their common name doesn’t just describe their scent, but also where they live.

At least three genera (Splachnum, Tetraplodon, and Tayloria) in the family Splachnaceae include species that go by the common name, dung moss. All Splachnum and Tetraplodon species and many species in the genus Tayloria are entomophilous. Entomophily is a “pollination syndrome”, a subject we will explore more thoroughly in future posts, in which pollen or spores are distributed by insects. Compare this to anemophily, or wind pollination, which is the more common way that moss spores are distributed. In fact, dung mosses are the only mosses known to exhibit entomophily.

Dung Moss (photo credit: wikimedia commons)

Dung Moss (photo credit: wikimedia commons)

Before we go too much further, it’s probably important to have a basic understanding of how mosses differ from other plants. Mosses are in a group of non-vascular and non-flowering plants called bryophytes. Vascular tissues are the means by which water and nutrients are transported to and from different plant parts. Lacking vascular tissues, water and nutrients are simply absorbed by the leaves of bryophytes (although some species have structures akin to vascular tissue), which is why they typically grow low to the ground and in moist environments. Bryophytes also lack true roots and instead have rhizoids, threadlike structures that anchor the plants to the ground or to some other substrate (such as dung).

Another major distinction between bryophytes and other plants is that bryophytes spend most of their life cycle as a haploid gametophyte rather than a diploid sporophyte (haploid meaning that it only has one set of chromosomes; diploid meaning that there are two sets of chromosomes, one from the father and one from the mother). In most plants, the haploid gametophyte is a sperm (pollen) or an egg. In bryophytes, the familiar green, leafy structure is actually the gametophyte. The gametophyte houses sperm and egg cells, and when the egg is fertilized by sperm it forms a zygote that develops into the sporophyte structure which extends above the leafy gametophyte. A capsule at the top of the sporophyte contains spores which are eventually released and, upon finding themselves on a suitable substrate in a hospitable environment, germinate to produce new plants. The spore then is comparable to a seed in vascular, seed-bearing plants.

photo credit: wikimedia commons

photo credit: wikimedia commons

As stated earlier, the spores of most mosses are distributed by wind. Dung mosses, on the other hand, employ flies in the distribution of their spores. They attract the flies by emitting scents that only flies can love from an area on the capsule of the sporophyte called the apophysis. This area is often enlarged and brightly colored in yellow, magenta, or red, giving it a flower-like appearance which acts as a visual attractant. The smells emitted vary depending on the type of substrate a particular species of dung moss has become adapted to living on. Some dung mosses grow on the dung of herbivores and others on the dung of carnivores. Some even prefer the dung of a particular group of animals; for example, a population of Tetraplodon fuegiensis was found to be restricted to the feces and remains of foxes. However, dung is not the only material that dung mosses call home.  Certain species grow on carrion, skeletal remains, or antlers. The smells these species produce attract flies that prefer dead flesh and bone in various states of decay.

Yellow Moosedung Moss (Splachnum luteum) has one of the largest and showiest sporophytes. (photo credit: www.eol.org)

Yellow Moosedung Moss (Splachnum luteum) has one of the largest and showiest sporophytes. (photo credit: www.eol.org)

The spores of dung mosses are small and sticky. When a fly visits these plants, the spores adhere to its body in clumps. The fly then moves on to its substrate of choice to lay its eggs, and the spores are deposited where they will then germinate and grow into new moss plants. Flies that visit dung mosses receive nothing in return for doing so, but instead are simply “tricked” into disseminating the propagules. The story is similar with corpse flowers and carrion flowers; flies are drawn in by the smells and recruited to transmit pollen but receive no nectar reward for their work.

There are 73 species in the Splachnaceae family, and nearly half of these species are dung mosses. These mosses are mostly found in temperate habitats in both the northern and southern hemispheres, with a few species occurring in the mountains of subtropical regions. They can be found in both wet and relatively dry habitats. Dung mosses are generally fast growing but short lived, with some lasting only about 2 years. It isn’t entirely clear how and why mosses in this family evolved to become entomophilous, but one major benefit of being this way is that their spores are reliably deposited on suitable habitat. Because of this directed dispersal, they can produce fewer and smaller spores, which is an economical use of resources.

Sporophytes of Splachnum vasculosum (photo credit: www.eol.org)

Sporophytes of Splachnum vasculosum (photo credit: www.eol.org)

References

Koponen, A. 2009. Entomophily in the Splachnaceae. Botanical Journal of the Linnean Society 104: 115-127.

Marino, P., R. Raguso, and B. Goffinet. 2009. The ecology and evolution of fly dispersed dung mosses (Family Splachnaceae): Manipulating insect behavior through odour and visual cues. Symbiosis 47: 61-76.

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My Carrion Flowers

In April of last year, a box of stem cuttings arrived in my mailbox. They were sent to me by a friend in Colorado called Sandra (you may know her from one of her many ventures: Greenwoman Magazine, Greenwoman Publishing, Flora’s Forum, etc.). Sandra’s carrion flower had bloomed that spring, a stinky but delightful occasion. In her excitement, she asked if I would be interested in growing some carrion flowers of my own. Not one to turn down the chance to try my hand at cultivating something unusual, I gladly accepted her offer of a few cuttings sent via Priority Mail. Six cuttings arrived shortly thereafter, and upon reading through some instructions on the internet, I nestled them into their new home and hoped they would put down roots and stay a while.

carrion flower cuttings

There are several species of plants that are referred to commonly as carrion flower. The plant parts I received from Sandra are in the genus Stapelia (family: Apocynaceae or dogbane family), also known commonly as African starfish flower. There are around 100 species in the genus Stapelia, and they all originate from tropical and southern Africa, mostly in arid regions.

Stapelias are short-lived, low-growing, perennial succulents. Their stems typically stand erect and are produced along stolons (above ground runners), creating a tight clump of stems that appear cactus-like. Each stem has 4-6 flattened vertical flanks, giving it a cross or star shape when looking down from the top. On the outside edges of the flanks are a series of rudimentary leaves protruding from tubercles (wart-like growths), giving the stems a spiny appearance. The stems are usually green but can also be red or mottled with red or purple.

The flowers of Stapelia are the real show. They are produced at or near the base of the stem and have a star-shaped corolla with five fused petals that come to sharp points. The corolla has a wrinkly look and is often hairy, especially along the margins. Flowers can be variations of red, brown, yellow, and purple. In some species they can reach up to 18 inches wide. It is a unique looking flower, but even more unique is its scent. Because Stapelia flowers are pollinated by flies, they emit the scent of rotting animal flesh, an odor that flies can truly appreciate. In fact, flies can be so deceived by the appearance and scent of the flowers that they occasionally lay their eggs on or near them, expecting them to be a food source for their emerging larva.

Stapelia variegate (photo credit: eol.org)

Stapelia variegata (photo credit: eol.org)

Stapelia is easily propagated, especially by stem cuttings. Allow cuttings to dry in a cool, shady location for 48 hours and then stick them in a well-drained potting soil mix. Water moderately (preferably from below by placing the container in a tray and then filling the tray with water). Cuttings should root easily. All six of mine did.

Keep Stapelia in a sunny or mostly sunny location. If you live in USDA hardiness zone 9 or above, you can grow Stapelia outdoors. Otherwise, keep it indoors near a window that gets lots of sun. The main thing you will have to worry about is stem rot due to over watering. Grow Stapelia in a well-drained soil mix, water from below, and allow soil to dry out between waterings in order to avoid this.

Stapelia variegata (photo credit: eol.org)

Stapelia variegata (photo credit: eol.org)

As for me and my carrion flowers, like I said earlier, all six cuttings rooted. I transplanted one of them. Of the five left in the original pot, one rotted a couple weeks ago and another rotted during the writing of this post. The remaining ones still look healthy, but none of them have grown much since they rooted. The main problem I am having is that my house does not let in much sunlight. What appears relatively bright to me is probably cave-like to my carrion flowers. Until I remedy that situation, they may not grow much, they could continue to rot, and they probably won’t flower any time soon. However, if anything changes and I do get a flower out of them, I will make it a point to let you know. And Sandra will be proud.

stapelia today_edit

 

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Corpse Flower Blooms Again

It is not often that a plant in bloom makes headlines, but that is precisely what happened last week when another corpse flower bloomed at Missouri Botanical Garden. Amorphophallus titanum, commonly known as titan arum or corpse flower, is a rare species, both in cultivation and in the wild. It also rarely flowers, and when it does, the bloom only lasts for a few short days. It has the largest known unbranched inflorescence, and its flowers give off the scent of rotting flesh. For all these reasons, it is understandable why a blooming corpse flower might make the news.

Titan arums naturally occur in the western portion of an Indonesian island called Sumatra. Their future is threatened because they occur in rainforests that are currently being deforested for timber and palm oil production. Deforestation is also threatening the survival of the rhinoceros hornbill, a bird that is an important seed distributor of titan arums. Today there are a few hundred titan arums in cultivation in botanical gardens throughout the world. They are a difficult species to cultivate, but their presence in botanical gardens is important in order to learn more about them and to help educate the public about conservation efforts.

Amorphophaulls titanium, titan arum (photo credit: eol.org)

(photo credit: eol.org)

Titan arums are in the arum family (Araceae), a family that consists of around 107 genera including Caladium (elephant ears), Arisaema (jack-in-the-pulpits), and Wolffia (duckweeds), a genus that wins the records for smallest flowering plant and smallest fruit. Titan arums are famous for their giant inflorescence, which can reach more than 10 feet tall. The flowering stalk is known botanically as a spadix, a fleshy stem in the shape of a spike that is covered with small flowers. The spadix of titan arums are wrapped with a leaf-like sheath called a spathe. Upon blooming, the temperature inside the spathe rises and the flowers begin to release a very foul odor, similar to the smell of rotting flesh. This attracts pollinating insects such as carrion beetles, sweat bees, and flesh flies, which get trapped inside the sheath and covered with pollen. After a few hours the top of the spadix begins to wither, allowing the insects to escape, off to pollinate a neighboring corpse flower [the spadix includes male and female flowers, which mature at different times in order to prevent self-pollination]. Once pollinated, the flowers begin to form small red fruits which are eaten by birds. The seeds are then dispersed in their droppings.

The large, stinky inflorescence is not the only structure that gives titan arums their fame. They are also known for their massive single leaf, which can reach up to 20 feet tall and 15 feet wide, the size of a large shrub or small tree. All of this growth is produced from an enormous underground storage organ called a corm. The corms of mature titan arums typically weigh more than 100 pounds, with some known to weigh more than 200 pounds. Titan arums bloom only after the corms have reached a mature size, which takes from seven to ten years. After that they bloom about once a year or once every other year, depending on when the corm has accumulated enough nutrients to support the giant flowering structure.

Below are two time lapse videos of titan arums in bloom. The first is from Missouri Botanical Garden, and the second is from United States Botanic Garden.

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