2015: Year in Review

Raise your glass. 2015 has come to a close, and Awkward Botany is turning three. Two great reasons to celebrate.

I started the year with the goal of posting at least once per week. Consider that goal accomplished, with a couple of bonus posts thrown in for good measure. I had also deemed 2015 the “Year of Pollination.” The underlying purpose was to teach myself more about pollinators and pollination while also sharing my interest in pollination biology with the wider world. That endeavor yielded 17 posts. There is still so much to learn, but we are making some headway. I started two new series of posts (Poisonous Plants and Botany in Popular Culture) and I continued with two others (Ethnobotany and Drought Tolerant Plants). I also went on a couple of field trips and wrote a few book reviews. All of that is reflected below in “Table of Contents” fashion.

Year of Pollination:

Botany in Popular Culture

Poisonous Plants

Ethnobotany

Drought Tolerant Plants

Book Reviews

Field Trips

Three posts that perhaps didn’t get the attention they deserve:

juniper in the snow

Going forward, I will continue to post regularly – as there is no shortage of plant-related things to write about – but I will likely take a week off here and there. I have other projects in mind – some related to Awkward Botany, some not – that will certainly demand much of my attention and time. I have some big ideas for Awkward Botany and beyond, and I will share those with the wide world in due time. For now, I would just like to say thanks all for reading, for commenting, and for sharing Awkward Botany with your friends. Overall, it has been a great year here at Awkward Botany headquarters, and I have you to thank for that. I feel privileged to be part of a community that is infatuated with plants and is fascinated by the natural world.

Good riddance to 2015. It was good, but it gets better. Now we look ahead to 2016. May it be filled with peace, love, and botany.

Year of Pollination: Botanical Terms for Pollination, part two

“The stage is set for reproduction when, by one means or another, compatible pollen comes to rest on a flower’s stigma. Of the two cells within a pollen grain, one is destined to grow into a long tube, a pollen tube, that penetrates the pistil’s tissues in search of a microscopic opening in one of the ovules, located in the ovary. … The second of a pollen grain’s cells divides to become two sperm that move through the pollen tube and enter the ovule.” – Brian Capon, Botany for Gardeners

“Once pollination occurs, the next step is fertilization. Pollen deposited on the sticky stigma generates a fine pollen tube that conveys the sperm through the style to the ovary, where the ovules, or eggs, have developed. After fertilization, the rest of the flower parts wither and are shed as the ovary swells with seed development.” – Rick Imes, The Practical Botanist

Pollination tells the story of a pollen grain leaving an anther by some means – be it wind, water, or animal – and finding itself deposited atop a stigma. As long as the pollen and stigma are compatible, the sex act proceeds. In other words, the pollen grain germinates. One of the pollen grain’s cells – the tube nucleus – grows down the length of the style, forming a tube through which two sperm nuclei can travel. The sperm nuclei enter the ovary and then, by way of a micropyle, enter an ovule. Inside the ovule is the female gametophyte (also referred to as the embryo sac). One sperm nucleus unites with the egg nucleus to form a zygote. The remaining sperm nucleus unites with two polar nuclei to form a triploid cell which becomes the endosperm. The sex act is complete.

The illustration on the left includes the cross-section of a pistil showing the inside the ovary where pollen tubes have made their way to the ovules. The illustration on the right shows pollen grains germinating on a stigma and their pollen tubes begining to work their way down the style. (photo credit: wikimedia commons)

The illustration on the left includes the cross section of a pistil showing the inside of the ovary where pollen tubes have made their way to the ovules. The illustration on the right shows pollen grains germinating on a stigma and pollen tubes as they work their way down the style. (image credit: wikimedia commons)

The zygote divides by mitosis to become an embryo. The endosperm nourishes the development of the embryo. The ovule matures into a seed, and the ovary develops into a fruit. During this process, the remaining parts of the flower wither and fall away. In some cases, certain flower parts remain attached to the fruit or become part of the fruit. The flesh of an apple, for example, is formed from the carpels and the receptacle (the thickened end of a flower stem – peduncle – to which the parts of a flower are attached).

As the seed matures, the endosperm is either used up or persists to help nourish the embryonic plant after germination. Mature seeds that are abundant in endosperm are called albuminous. Examples include wheat, corn, and other grasses and grains. Mature seeds with endosperm that is either highly reduced or absent are called exalbuminous – beans and peas, for example. Certain species – like orchids – do not produce endosperm at all.

The cross section of a corn kernel showing the endosperm and the embryo (image credit: Encyclopedia Britannica Kids)

The cross section of a corn kernel showing the endosperm and the embryo (image credit: Encyclopedia Britannica Kids)

It is fascinating to consider that virtually every seed we encounter is the result of a single pollen grain making its way from an anther to a stigma, growing a narrow tube down a style, and fertilizing a single ovule. [Of course there are always exceptions. Some plants can produce seeds asexually. See apomixis.] Think of this the next time you are eating corn on the cob or popcorn – each kernel is a single seed – or slicing open a pomegranate to reveal the hundreds of juicy seeds inside. Or better yet, when you are eating the flesh or drinking the milk of a coconut. You are enjoying the solid and liquid endosperm of one very large seed.

Much more can be said about pollination and the events surrounding it, but we’ll save that for future posts. The “Year of Pollination” may be coming to an end, but there remains much to discover and report concerning the subject. For now, here is a fun video to help us review what we’ve learned so far:

 

Also, take a look at this TED talk: The Hidden Beauty of Pollination by Louie Schwartzberg

And finally, just as the “Year of Pollination” was coming to an end I was introduced to a superb blog called The Amateur Anthecologist. Not only did it teach me that “anthecology” is a term synonymous with pollination biology, it has a great series of posts called “A Year of Pollinators” that showcases photographs and information that the author has collected for various groups of pollinators over the past year. The series includes posts about Bees, Wasps, Moths and ButterfliesFlies, and Beetles, Bugs, and Spiders.

Year of Pollination: Botanical Terms for Pollination, part one

When I began this series of posts, I didn’t have a clear vision of what it would be. I had a budding interest in pollination biology and was anxious to learn all that I could. I figured that calling 2015 the “Year of Pollination” and writing a bunch of pollination-themed posts would help me do that. And it has. However, now that the year is coming to a close, I realize that I neglected to start at the beginning. Typical me.

What is pollination? Why does it matter? The answers to these questions seemed pretty obvious; so obvious, in fact, that I didn’t even think to ask them. That being said, for these last two “Year of Pollination” posts (and the final posts of the year), I am going back to the basics by defining pollination and exploring some of the terms associated with it. One thing is certain, there is still much to be discovered in the field of pollination biology. Making those discoveries starts with a solid understanding of the basics.

Pollination simply defined is the transfer of pollen from an anther to a stigma or – in gymnosperms – from a male cone to a female cone. Essentially, it is one aspect of plant sex, albeit a very important one. Sexual reproduction is one way that plants multiply. Many plants can also reproduce asexually. Asexual reproduction typically requires less energy and resources – no need for flowers, pollen, nectar, seeds, fruit, etc. – and can be accomplished by a single individual without any outside help; however, there is no gene mixing (asexually reproduced offspring are clones) and dispersal is limited (consider the “runners” on a strawberry plant producing plantlets adjacent to the mother plant).

To simplify things, we will consider only pollination that occurs among angiosperms (flowering plants); pollination/plant sex in gymnosperms will be discussed at another time. Despite angiosperms being the youngest group of plants evolutionarily speaking, it is the largest group and thus the type we encounter most.

A flower is a modified shoot and the reproductive structure of a flowering plant. Flowers are made up of a number of parts, the two most important being the reproductive organs. The androecium is a collective term for the stamens (what we consider the male sex organs). A stamen is composed of a filament (or stalk) topped with an anther – where pollen (plant sperm) is produced. The gynoecium is the collective term for the pistil (what we consider the female sex organ). This organ is also referred to as a carpel or carpels; this quick guide helps sort that out. A pistil consists of the ovary (which contains the ovules), and a style (or stalk) topped with a stigma – where pollen is deposited. In some cases, flowers have both male and female reproductive organs. In other cases, they have one or the other.

photo credit: wikimedia commons

photo credit: wikimedia commons

When pollen is moved from an anther of one plant to a stigma of another plant, cross-pollination has occurred. When pollen is moved from an anther of one plant to a stigma of the same plant, self-pollination has occurred. Cross-pollination allows for gene transfer, and thus novel genotypes. Self-pollination is akin to asexual production in that offspring are practically identical to the parent. However, where pollinators are limited or where plant populations are small and there is little chance for cross-pollination, self-pollination enables reproduction.

Many species of plants are unable to self-pollinate. In fact, plants have evolved strategies to ensure cross-pollination. In some cases, the stamens and pistils mature at different times so that when pollen is released the stigmas are not ready to receive it or, conversely, the stigmas are receptive before the pollen has been released. In other cases, stigmas are able to recognize their own pollen and will reject it or inhibit it from germinating. Other strategies include producing flowers with stamens and pistils that differ dramatically in size so as to discourage pollen transfer, producing separate male and female flowers on the same plant (monoecy), and producing separate male and female flowers on different plants (dioecy).

As stated earlier, the essence of pollination is getting the pollen from the anthers to the stigmas. Reproduction is an expensive process, so ensuring that this sex act takes place is vital. This is the reason why flowers are often showy, colorful, and fragrant. However, many plants rely on the wind to aid them in pollination (anemophily), and so their flowers are small, inconspicuous, and lack certain parts. They produce massive amounts of tiny, light-weight pollen grains, many of which never reach their intended destination. Grasses, rushes, sedges, and reeds are pollinated this way, as well as many trees (elms, oaks, birches, etc.) Some aquatic plants transport their pollen from anther to stigma via water (hydrophily), and their flowers are also simple, diminutive, and produce loads of pollen.

Inforescence of big bluestem (Andropogon gerardii), a wind pollinated plant - pohto credit: wikimedia commons

Inflorescence of big bluestem (Andropogon gerardii), a wind pollinated plant – photo credit: wikimedia commons

Plants that employ animals as pollinators tend to have flowers that we find the most attractive and interesting. They come in all shapes, sizes, and colors and are anywhere from odorless to highly fragrant. Odors vary from sweet to bitter to foul. Many flowers offer nectar as a reward for a pollinator’s service. The nectar is produced in special glands called nectaries deep within the flowers, inviting pollinators to enter the flower where they can be dusted with pollen. The reward is often advertised using nectar guides – patterns of darker colors inside the corolla that direct pollinators towards the nectar. Some of these nectar guides are composed of pigments that reflect the sun’s ultraviolet light – they are invisible to humans but are a sight to behold for many insects.

In part two, we will learn what happens once the pollen has reached the stigma – post-pollination, in other words. But first, a little more about pollen. The term pollen actually refers to a collection of pollen grains. Here is how Michael Allaby defines “pollen grain” in his book The Dictionary of Science for Gardeners: “In seed plants, a structure produced in a microsporangium that contains one tube nucleus and two sperm nuclei, all of them haploid, enclosed by an inner wall rich in cellulose and a very tough outer wall made mainly from sporopollenin. A pollen grain is a gametophyte.”

A pollen grain’s tough outer wall is called exine, and this is what Allaby has to say about that: “It resists decay, and the overall shape of the grain and its surface markings are characteristic for a plant family, sometimes for a genus or even a species. Study of pollen grains preserved in sedimentary deposits, called palynology or pollen analysis, makes it possible to reconstruct past plant communities and, therefore, environments.”

Scanning electron microscope image of pollen grains from narrowleaf evening primrose (Oenothera fruticosa) - photo credit: wikimedia commons

Scanning electron microscope image of pollen grains from narrowleaf evening primrose (Oenothera fruticosa) – photo credit: wikimedia commons

Year of Pollination: Scarlet Gilia and Its Pollinators

Flowers that are visited and/or pollinated by hummingbirds typically fit the following description: petals are brightly colored, often red; petals are fused to form a long, narrow tube; a “landing pad” is absent; abundant nectar is produced deep within the flower; and fragrance is weak or nonexistent. Scarlet gilia (Ipomopsis aggregata) is a typical example of such a flower, and hummingbirds are indeed among its most common visitors. But there is so much more to the story.

Scarlet gilia (also commonly known as skyrocket) is a wildflower in the phlox family (Polemoniaceae) that occurs in many parts of western North America. It is considered a biennial or short-lived perennial. It spends the first year or so of its life as a compact rosette of fern-like leaves. Later it sends up a branched, flowering stem that can reach 5 feet tall or more. The flowers are slender, trumpet-shaped, and composed of five fused petals that flare outward creating five prominent, pointed lobes. They are self-incompatible and require a pollinator in order to set seed. The stamens of an individual flower produce mature pollen before the stigma of that flower is ready to receive it – this is called protandry and is one mechanism of self-incompatibility.

The rosette of scarlet gilia (Ipomopsis aggregata)

The rosette of scarlet gilia (Ipomopsis aggregata)

The flowering period of scarlet gilia can last several months. Depending on the location, it can begin in mid-summer and continue through the fall. During this period, it produces dozens of flowers. It is also at this time that it runs the risk of being browsed by elk, mule deer, and other animals. This doesn’t necessarily set it back though, as it has the potential to respond by producing additional flowering stalks and more flowers. Its flowers are visited by a variety of pollinators including bumblebees, hawkmoths, butterflies, syrphid flies, solitary bees, and of course, hummingbirds. But hummingbirds, in many parts of scarlet gilia’s range are migratory, and that’s where things get interesting.

Early flowers of scarlet gilia are usually red. As the season progresses, flowers slowly shift from red to pink. In some cases, they lose all pigmentation and become white. In the early 1980’s, pollination biologists Ken Paige and Thomas Whitman set out to determine the reason for this shift in flower color. They spent three years observing a population of scarlet gilia on Fern Mountain near Flagstaff, Arizona. They noted that the change in flower color corresponded with the migration of hummingbirds and that the now lighter colored flowers continued to be pollinated by hawkmoths until the end of the flowering season.

ipomopsis aggregata

A series of experiments and observations led them to conclude that hummingbirds prefer darker colored flowers and hawkmoths prefer lighter colored flowers. By shifting to a lighter flower color, scarlet gilia appeared to be taking advantage of remaining pollinators after hummingbirds had migrated. They also concluded that the color change was not the cause of hummingbird migration since other flowers with nectar-rich, red, tubular flowers (specifically Penstemon barbatus) remained available in the area throughout their migration. It was also noted that the flowers of scarlet gilia shifted the timing of nectar production, presumably to better match the behavior of hawkmoths which are more active in the evenings.

No plants were observed shifting from light colored flowers to dark colored flowers, which further supported their conclusion. They also compared the population they studied to populations that do not lose their hummingbird pollinator and noted that when hummingbirds remain, the flowers of scarlet gilia don’t change color.

Scarlet gilia (Ipomopsis aggregata) with white flowers

Scarlet gilia (Ipomopsis aggregata) with white flowers

But just how effective are hummingbirds as pollinators of scarlet gilia? A seperate study carried out by a different group of researchers determined that, while hummingbirds were “the most common floral visitor,” long-tongued bumblebees were the more effective pollinator when it came to pollen deposition and seed set. The study involved observations of a scarlet gilia population in Colorado over a 5 year period. Considering how well the floral traits of scarlet gilia match up with the hummingbird pollination syndrome, it is surprising to learn that long-tongued bumblebees are comparatively more effective at pollinating them.

This study provides further evidence against strict adherence to pollination syndromes and the most effective pollinator principle, both of which imply specialized plant-pollinator interactions. (I wrote about these topics here, here, and here in earlier Year of Pollination posts.) In their discussion, the authors propose two possible explanations as to why scarlet gilia, despite its phenotypic floral traits, does not appear to be specialized. One explanation is that “natural selection favors a specialized [floral] morphology that excludes all but a single type of visitor, but there are constraints on achieving this outcome.” Perhaps the pollinators aren’t cooperating; their opportunism is leading them to “exploit flowers on which they can realize an energetic profit, even if they do not mechanically ‘fit’ very well.” The “sensory abilities” of the pollinators may be “broadly tuned,” making it difficult for plants to develop flowers with “private signals detectable only by specific types of pollinators.”

The second explanation proposed by the authors is that “selection favors some degree of floral generalization, but that flowers can retain features that adapt them to a particular type of pollinator in spite of generalization.” In the case of scarlet gilia, specialization could be detrimental because after they send up their flower stalks, they are doomed to die. This gives them only one season to set seed, and if hummingbirds are either not available that year or only available in limited numbers, a scarlet gilia population can lose the opportunity to reproduce. As the authors put it, “the fact that individual plants enjoy only a single season of reproduction, suggests the value of ‘backup’ pollinators.” This may also explain why flower color shifts in order to take full advantage of hawkmoth pollination after hummingbirds are gone.

Scarlet gilia is not only a beautiful and widespread wildflower, but also a plant with a very interesting story. Follow the links below to learn more about this fascinating plant:

Year of Pollination: Hand Pollinating Cucurbits

Because of their large, open, unisexual flowers, plants in the gourd family are perfect for practicing hand pollination. There are several species in this family that are commonly grown in gardens, and all can be hand pollinated. Hand pollination of cucurbits is most often done when there are problems with pollination (lack of pollinators, etc.) or for seed saving purposes (i.e. to ensure that a variety breeds true). It can also be done just for fun, and that’s mostly what this post is about.

But first, if your goal is to save seeds and maintain the integrity of the varieties you are growing, there are a few things to keep in mind. Cucumbers, melons, and watermelons are all different species (Cucumis sativus, Cucumis melo, and Citrullus lanatus respectively), so you won’t have to worry about crosses between these crops. You will, however, have to worry about crosses between different varieties within individual species. So, for example, if you are growing multiple varieties of cucumbers – or if your close neighbors are also growing cucumbers – you should hand pollinate. Summer squash, winter squash, pumpkins, and some gourds are members of at least four species in the genus Cucurbita (C. pepo, C. maxima, C. mixta, and C. moschata). There is a possibility of hybridization between some of these species as well as between varieties within the same species, so precautions should definitely be taken when saving seeds for these crops. This can mean, along with hand pollination, placing bags over flowers so that bees are unable to bring in pollen from “the wrong” plants.

There are plenty of great resources about saving seeds that offer much more detail than I have gone into here, one of which is a book by Marc Rogers called Saving Seeds. Consult such resources if you would like to try your hand at seed saving. It’s easier than you might think, and it’s very rewarding.

Regardless why you are hand pollinating your cucurbits, the first step in the process is differentiating a male flower from a female flower. This is simple. Female flowers in the family Cucurbitaceae have inferior ovaries, meaning that the ovary sits below the area where the petals and other flower parts are attached. The ovaries are quite pronounced and resemble a miniature fruit. The male flowers lack ovaries, so instead are simply attached to a slender stem. You can also observe the sex organs themselves – male flowers have stamens, female flowers have carpels. Male and female flowers may also be located on different areas of the plant and may open at different times of the day. All that being said, the most obvious indication is the “mini-fruit” at the base of the flower or lack thereof.

Cucurbit flowers: male (top) and female (bottom) - photo credit: wikimedia commons

Cucurbit flowers: male (top two photos) and female (bottom two photos) – image credit: wikimedia commons

Once you have identified your flowers, you have a limited amount of time to hand pollinate them. It’s best to find flowers that are just starting to open, as the female flowers may only be receptive for as little as 24 hours. You can use a cotton swab to gather pollen from the male flower, or you can simply pluck the flower from the plant, remove the petals, and touch the pollen-loaded anthers to the stigmas of a female flower. Either way, you must get the pollen from the male parts of a flower to the female parts of a flower as that is the essence of pollination. Simply put, it’s plant sex. Play some soft jazz while you do it if you want to.

A honeybee in a squash flower

A female squash flower with honeybee inside

A honeybee covered in pollen drinking the nectar of a female squash flower

Honeybee covered in pollen drinking the nectar of a female squash flower

As with saving seeds, there are a lot of resources out there explaining the details of hand pollinating cucurbit flowers, including this guide from Missouri Botanical Garden and the following You Tube Video.

 

While we are on the subject of cucurbit flowers, it should be noted that squash flowers are edible and can be prepared in a variety of ways, as described in this post at The Kitchn. Just another reason to be impressed by this amazing group of plants.

Year of Pollination: Bumblebees and Climate Change

Bumblebees, generally speaking, are having a rough time. In a world increasingly dominated by humans, some bumblebee species continue to thrive while many others are seriously struggling. Several are nearing extinction. A recent study involving 67 species of European and North American bumblebees concluded that climate change is having a major impact. Bumblebees do not appear to be migrating north in response to warming climates – a hesitation that could spell disaster.

There are over 250 species of bumblebees worldwide (46 are found in North America north of Mexico). Unlike other bees, whose diversity is greatest in Mediterranean climates, bumblebee diversity is highest in cool, temperate climates and montane regions. The majority of bumblebee species are native to the Northern Hemisphere; a few species are native to South America, and a handful of species from Europe have been introduced to New Zealand and Tasmania. Some species of bumblebees, such as the polar bumble bee (Bombus polaris) and the forest bumble bee (Bombus sylvicola), can be found in extreme cold climates and are among a select group of pollinators found in such areas.

The field guide, Bumble Bees of North America, by Paul Williams, et al. provides this description:

“Bumble bees are very hairy bees with combinations of contrasting bright colors, mostly black and yellow, sometimes with various combinations of red or white. They have two pairs of wings that are usually folded back over the abdomen while they are foraging on flowers, or hooked together as a single unit when in flight. Bumble bees also have slender elbowed antennae, and females of the pollen-collecting species have the hind tibia expanded, slightly concave, and fringed with long hairs to form a pollen basket or corbicula.”

Most bee species are solitary insects; bumblebees, like honeybees, are social insects. Unlike honeybees, bumblebee colonies begin with a new queen each year. New queens, after mating in late summer, overwinter in a protected area and emerge in the spring. They then search for food and a nesting site. Suitable nests include abandoned rodent dens,  the bases of bunchgrasses, hollow logs, and human-made structures. They build up a colony of workers which maintain the nest and forage for food and other resources. As the season comes to a close, the queen produces males and new queens. The new queens mate, go into hibernation, and the rest of the bumblebee colony dies off.

Brown-belted Bumblebee (Bombus griseocollis) - photo credit: wikimedia commons

Brown-belted Bumblebee (Bombus griseocollis) – photo credit: wikimedia commons

Bumblebees face numerous threats, both natural and human-caused. Despite their defensive sting, they are regularly eaten or attacked by various mammals, birds, and invertebrates. They are also host to a variety of pests, parasites, and pathogens, some of which have been introduced or exacerbated by human activities. The commercial bee industry is particularly at fault for the spread of certain maladies. Other major threats include loss of habitat and excessive and/or poorly timed use of insecticides. One looming threat that new research suggests is especially concerning is climate change.

A group of researchers from various institutions looked at the historical ranges of 67 species of bumblebees in Europe and North America over a 110 year period. They “measured differences in species’ northern and southern range limits, the warmest or coolest temperatures occupied, and their mean elevations in three periods relative to a baseline period.” They found that on both continents bumblebees are not tracking climate change by expanding their northern range limits and that their southern range limits are shrinking. They also observed that within the southern range limits, some bumblebee species have retreated to higher elevations. They investigated land use changes and pesticide applications (in the US only) to determine the effect they had on the results. While these things certainly affect populations on an individual level, climate change was determined to be the most important factor that lead to nearly universal range contractions of the bumblebees in this study.

The question then is why are they not tracking changing climates the same way that many other species of plants and animals have already been observed doing? Bumblebees evolved in cooler climates, so shrinking southern range limits is not as surprising as the bumblebees’ delay in moving north. Many factors may be contributing to this phenomenon including lack of specialized habitats beyond their historical ranges, daylength differences, and population dynamics. The researchers call for further investigation in order to better evaluate this observed “range compression.” They also suggest experimenting with assisted migration of certain bumblebee colonies, which in general is a controversial topic among conservation biologists. (Read more about this study here.)

Buff-tailed Bumblebee (Bombus terrestris) - photo credit: wikimedia commons

Buff-tailed Bumblebee (Bombus terrestris) – photo credit: wikimedia commons

The loss of bumblebees is concerning because they play a prominent role in the various ecosystems in which they live. They are prolific and highly effective pollinators of both agricultural crops and native plants, and they are also a major component in the food web. Some species of plants “prefer” the pollination services of bumblebees, such as those in the family Solanaceae. Many plants in this family benefit greatly from buzz pollination – a process in which a bumblebee (or occasionally bees of other species) grabs hold of the flower and vibrates its body, dislodging the pollen.

Participating in bumblebee conservation is simple. It’s similar to any other kind of pollinator conservation. Just learning about the pollinators in your region and being mindful of them can make a big difference. If you own or rent property and have space for a garden (even if its just a few containters on a patio), choose plants that provide food for bumblebees, including spring and summer bloomers. If you live in North America, this Xerces Society publication and the field guide mentioned above are great resources that can help you determine which plants are best for your region. Additionally, if you are working in your yard and happen upon a hibernating queen or a bumblebee nest, do your best not to disturb it. It may disrupt your gardening plans for a season, but the bumblebee sightings and the pollination service they provide will be worth it.

One family of plants in particular that you should consider representing in your yard is the legume family (Fabaceae). Bumblebees are commonly seen pollinating plants in this family, and because these plants have the ability to convert nitrogen in the air into fertilizer, their pollen is especially rich in protein. In his book, A Sting in the Tale, Dave Goulson describes the relationship between bumblebees and legumes:

“From a bumblebee’s perspective, legumes are among the most vital components of a wildflower meadow. Plants of this family include clovers, trefoils and vetches, as well as garden vegetables such as peas and beans, and they have an unusual trick that allows them to thrive in low-fertility soils. Their roots have nodules, small lumps inside which live Rhizobium, bacteria that can trap nitrogen from the air and turn it into a form usable by plants. … This relationship gave legumes a huge advantage in the days before artificial fertilizers were widely deployed. Ancient hay meadows are full of clovers, trefoils, vetches, meddicks and melilots, able to outcompete grasses because they alone have access to plentiful nutrients. Most of these plants are pollinated by bumblebees.”

More information about bumblebees and bumblebee conservation:

Bumblebee Conservation Trust

Bumble Bee Watch

BugGuide (Bombus)

The Xerces Society – Project Bumble Bee

Year of Pollination: Figs and Fig Wasps

This post originally appeared on Awkward Botany in November 2013. I’m reposting an updated version for the Year of Pollination series because it describes a very unique and incredibly interesting interaction between plant and pollinator. 

Ficus is a genus of plants in the family Moraceae that consists of trees, shrubs, and vines. Plants in this genus are commonly referred to as figs, and there are nearly 850 described species of them. The majority of fig species are found in tropical regions, however several occur in temperate regions as well. The domesticated fig (Ficus carica), also known as common fig, is widely cultivated throughout the world for its fruit.

common fig

Common Fig (Ficus carica) – photo credit: wikimedia commons

The fruit of figs, also called a fig, is considered a multiple fruit because it is formed from a cluster of flowers. A small fruit develops from each flower in the cluster, but they all grow together to form what appears to be a single fruit. The story becomes bizarre when you consider the location of the fig flowers. They are contained inside a structure called a syconium, which is essentially a modified fleshy stem. The syconium looks like an immature fig. Because they are completely enclosed inside syconia, the flowers are not visible from the outside, yet they must be pollinated in order to produce seeds and mature fruits.

This is where the fig wasps come in. “Fig wasp” is a term that refers to all species of chalcid wasps that breed exclusively inside of figs. Fig wasps are in the order Hymenoptera (superfamily Chalcidoidea) and represent at least five families of insects. Figs and fig wasps have coevolved over tens of millions of years, meaning that each species of fig could potentially have a specific species of fig wasp with which it has developed a mutualistic relationship. However, pollinator host sharing and host switching occurs frequently.

Fig wasps are tiny, mere millimeters in length, so they are not the same sort of wasps that you’ll find buzzing around you during your summer picnic. Fig wasps have to be small though, because in order to pollinate fig flowers they must find their way into a fig. Fortunately, there is a small opening at the base of the fig called an ostiole that has been adapted just for them.

What follows is a very basic description of the interaction between fig and fig wasp; due to the incredible diversity of figs and fig wasps, the specifics of the interactions are equally diverse.

First, a female wasp carrying the pollen of a fig from which she has recently emerged discovers a syconium that is ready to be pollinated. She finds the ostiole and begins to enter. She is tiny, but so is the opening, and so her wings, antennae, and/or legs can be ripped off in the process. No worries though, since she won’t be needing them anymore. Inside the syconium, she begins to lay her eggs inside the flowers. In doing so, the pollen she is carrying is rubbed off onto the stigmas of the flowers. After all her eggs are laid, the female wasp dies. The fig wasp larvae develop inside galls in the ovaries of the fig flowers, and they emerge from the galls once they have matured into adults. The adult males mate with the females and then begin the arduous task of chewing through the wall of the fig in order to let the females out. After completing this task, they die. The females then leave the figs, bringing pollen with them, and search for a fig of their own to enter and lay eggs. And the cycle continues.

But there is so much more to the story. For example, there are non-pollinating fig wasps that breed inside of figs but do not assist in pollination – freeloaders essentially. The story also differs if the species is monoecious (male and female flowers on the same plant) compared to dioecious (male and female flowers on different plants). It’s too much to cover here, but figweb.org is a great resource for fig and fig wasp information. Also check out the PBS documentary, The Queen of Trees.

 

 

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.

Year of Pollination: Mosquitoes as Pollinators

It is difficult to have positive feelings about mosquitoes, especially during summer months when they are out in droves and our exposed skin – soft, supple, and largely hair-free – is irresistible to them. We are viewed as walking blood meals by female mosquitoes who are simply trying to produce young – to perpetuate their species just like any other species endeavors to do. Unfortunately, we are left with small, annoying bumps in our skin – red, itchy, and painful – risking the possibility that the mosquitoes that just drew our blood may have passed along any number of mosquito-borne diseases, some (such as malaria) that potentially kill millions of people every year. For this, it is okay to hate mosquitoes and to long for the day of their complete eradication from the planet. However, their ecological roles (and yes, they do have some) are also worth considering.

There are more than 3,500 species of mosquito. Luckily, only 200 or so consume human blood. Mosquitoes go back at least 100 million years and have co-evolved with species of plants and animals found in diverse habitats around the world. Adult mosquitoes and their larvae (which live in standing water) provide food for a wide variety of creatures including birds, bats, insects, spiders, fish, frogs, lizards, and salamanders. Mosquito larvae also help break down organic matter in the bodies of water they inhabit. They even play an important role in the food webs found inside the pitchers of northern pitcher plants (Sarracenia spp.). Interestingly enough, Arctic mosquitoes influence the migration patterns of caribou. They emerge in swarms so big and so voracious that they have been said to kill caribou through either blood loss or asphyxiation.

However, blood is not the main food source of mosquitoes; flower nectar is. Males don’t consume blood at all, and females only consume it when they are producing eggs. Any insect that visits flowers for nectar has the potential to unwittingly collect pollen and transfer it to a nearby flower, thereby aiding in pollination. Mosquitoes are no exception. They have been observed acting as pollinators for a handful of species, and could be acting as pollinators for many more.

Bluntleaved orchid (Platanthera obtusata) is pollinated by mosquitoes. phot credit: wikimedia commons

Bluntleaved orchid (Platanthera obtusata) is pollinated by mosquitoes. photo credit: wikimedia commons

The scientific literature describes the pollination by mosquitoes of at least two plant species: Platanthera obtusata (syn. Habenaria obtusata) and Silene otites. P. obtusata – bluntleaved orchid – is found in cold, wet regions in North America and northern Eurasia. It is pollinated by mosquitoes from multiple genera including several species in the genus Aedes. Mosquitoes visit the flowers to feed on the nectar and, subsequently, pollinia (clusters of pollen) become attached to their eyes and are moved from flower to flower. This scenario likely plays out in other species of Arctic orchids as well*.

S. otites – Spanish catchfly – is a European species that is pollinated by mosquitoes and moths. Researches have been studying the floral odors of S. otites that attract mosquitoes, suggesting that determining the compounds involved in these odors “might lead to the development of new means of pest control and mosquito attractants and repellents.”

Northern House Mosquito (Culex pipiens) - one of the species of mosquitoes that has been observed pollinating Silene otitis. photo credit: www.eol.org

Northern House Mosquito (Culex pipiens) – one of the species of mosquitoes that has been observed pollinating Silene otites. photo credit: www.eol.org

Despite the list of functions that mosquitoes serve in their varied habitats, an article that appeared in Nature back in 2010 argues for wiping mosquitoes off the Earth, stating that “the ecological scar left by a missing mosquito would heal quickly as the niche was filled by other organisms.” And even though “thousands of plant species would lose a group of pollinators,” mosquitoes are not important pollinators of the “crops on which humans depend,” nor do they appear to be the sole pollinator of any single plant species [the species mentioned above are pollinated by other insects as well]. Eliminating mosquitoes, however, is more of a pipe dream than a realistic possibility as our “best efforts can’t seriously threaten an insect with few redeeming features.”

*Speaking of orchids and pollination, endless posts could be written about this incredibly fascinating and diverse group of plants and their equally fascinating and complex mechanisms surrounding pollination. There will be more to come on such topics. Meanwhile, it should be noted that orchids are also a notoriously threatened group of plants. To learn more about orchids and orchid conservation in North America, visit North American Orchid Conservation Center.

Read more about mosquito pollination here.

And now for your listening pleasure:

Year of Pollination: An Argentinian Cactus and Its Unlikely Pollinator

A few weeks ago I wrote about pollination syndromes – sets of floral triats that are said to attract specific groups of pollinators. In that post I discussed how pollination syndromes have largely fallen out of favor as a reliable method of predicting the pollinators that will visit particular flowers. In this post I review a recent study involving a species of cactus in Argentina that, as the authors state in their abstract, “adds another example to the growing body of mismatches between floral syndrome and observed pollinator.”

Denmoza rhodacantha is one of many species of cacti found in Argentina. It is the only species in its genus, and it is widely distributed across the east slopes and foothills of the Andes. It is a slow growing cactus, maintaining a globulous (globe-shaped) form through its juvenile phase and developing a columnar form as it reaches maturity. D. rhodacantha can reach up to 4 meters tall and can live beyond 100 years of age. Individual plants can begin flowering in their juvenile stage. Flowers are red, nectar rich, scentless, and tubular. The stigma is lobed and is surrounded by a dense grouping of stamens. Both male and female reproductive organs are extended above the corolla. The flowers have been described by multiple sources as being hummingbird pollinated, not based on direct observation of hummingbirds visiting the flowers, but rather due to the floral traits of the species.

Denmoza rhodacantha illustration - image credit: www.eol.org

Denmoza rhodacantha illustration  (image credit: www.eol.org)

In a paper entitled, Flowering phenology and observations on the pollination biology of South American cacti – Denmoza rhodacantha, which was published in volume 20 of Haseltonia (the yearbook of the Cactus and Succulent Society of America), Urs Eggli and Mario Giorgetta discuss their findings after making detailed observations of a population of D. rhodacantha in early 2013 and late 2013 – early 2014. The population consisted of about 30 individuals (both juveniles and adults) located in the Calchaqui Valley near the village of Angastaco, Argentina. At least three other species with “hummingbird-syndrome flowers” were noted in the area, and three species of hummingbirds were observed during the study periods. Over 100 observation hours were logged, and during that time “the studied plants, their flowering phenology, and flower and fruit visitors were documented by means of photographs and video.”

The flowers of D. rhodacantha only persist for a few short days, and in that time their sexual organs are only receptive for about 24 hours. The flowers are self-sterile and so require a pollinator to cross pollinate them. Despite their red, tubular shape and abundant nectar, no hummingbirds were observed visiting the flowers. One individual hummingbird approached but quickly turned away. Hummingbirds were, however, observed visiting the flowers of an associated species, Tecoma fulva ssp. garrocha. Instead, a species of halictid bee (possibly in the genus Dialictus) was regularly observed visiting the flowers of D. rhodacantha. The bees collected pollen on their hind legs and abdomen and were seen crawling across the lobes of the stigma. None of them were found feeding on the nectar. In one observation, a flower was visited by a bee that was “already heavily loaded with the typical violet-coloured pollen of Denmoza,” suggesting that this particular bee species was seeking out these flowers for their pollen. Small, unidentified beetles and ants were seen entering the flowers to consume nectar, however they didn’t appear to be capable of offering a pollination service.

D. rhodacantha populations have been observed in many cases to produce few fruits, suggesting that pollination success is minimal. The authors witnessed “very low fruit set” in the population that they were studying, which was “in marked contrast to the almost 100% fruit set rates of the sympatric cactus species at the study site.” This observation wasn’t of great concern to the authors though, because juvenile plants are present in observed populations, so recruitment appears to be occurring. In this study, dehisced fruits were “rapidly visited by several unidentified species of ants of different sizes.” The “scant pulp” was harvested by smaller ants, and larger ants carried away the seeds after “cleaning them from adhering pulp.”

The authors propose at least two reasons why hummingbirds avoid the flowers of D. rhodacantha. The first being that the native hummingbirds have bills that are too short to reach the nectar inside the long tubular flowers, and often the flowers barely extend beyond the spines of the cactus which may deter the hummingbirds from approaching. The second reason is that other plants in the area flower during the same period and have nectar that is easier to gather. The authors acknowledge that this is just speculation, but it could help explain why the flowers are pollinated instead by an insect (the opportunist, generalist halictid bee species) for whom the flowers “could be considered to be ill adapted.” The authors go on to say, “it should be kept in mind, however, that adaptions do not have to be perfect, as long as they work sufficiently well.”

Patagona gigas (giant hummingbird) was observed approaching the flower of a Denmoza rhodacantha but quickly turned away (photo credit: www.eol.org)

Patagona gigas (giant hummingbird) was observed approaching the flower of a Denmoza rhodacantha but quickly turned away (photo credit: www.eol.org)

More Year of Pollination posts on Awkward Botany: