September 30, 2015 by awkwardbotany

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 (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.

September 2, 2015 by awkwardbotany

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.

July 8, 2015 by awkwardbotany

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:

May 27, 2015 by awkwardbotany

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)

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.

Denmoza rhodacantha at Univeristy of California Botanical Garden at Berkeley (photo credit: wikimedia commons)

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)

Year of Pollination: More than Honey, etc.

When I decided to spend a year writing about pollinators and pollination, I specifically wanted to focus on pollinators besides the honey bee. Honey bees already get lots of attention, and there are loads of other pollinating organisms that are equally fascinating. But that’s just the thing, honey bees are incredibly fascinating. They have a strict and complex social structure, and they make honey – two things that have led humans to develop a strong relationship with them. We have been managing honey bees and exploiting their services for thousands of years, and we have spread them across the planet, bringing them with us wherever we go. In North America, honey bees are used to pollinate a significant portion of our pollinator-dependent crops, despite the fact that they are not native to this continent. In that sense, they are just another domesticated animal, artificially selected for our benefit.

It’s common knowledge that honey bees (and pollinators in general) have been having a rough time lately. Loss of habitat, urbanization, industrial farming practices, abundant pesticide use, and a variety of pests and diseases have been making life difficult for pollinators. Generally, when the plight of pollinators comes up in the news, reference is made to honey bees (or another charismatic pollinator, the monarch butterfly). News like this encourages people to take action. On the positive side, efforts made to protect honey bees can have the side benefit of protecting native pollinators since many of their needs are the same. On the negative side, evidence suggests that honey bees can compete with native pollinators for limited resources and can pass along pests and diseases. Swords are often double-edged, and there is no silver bullet.

In a recent conversation with a budding beekeeper, I was recommended the documentary, More than Honey. I decided to watch it, write a post about it, and call that the honey bee portion of the Year of Pollination. Part way through the movie, another documentary, Vanishing of the Bees, was recommended to me, and so I decided to watch both. Below are some thoughts about each film.

More than Honey

Written and directed by Swiss documentary filmmaker, Markus Imhoof, this beautifully shot, excellently narrated, meandering documentary thrusts viewers into incredibly intimate encounters with honey bees. Cameras follow bees on their flights and into their hives and get up close and personal footage of their daily lives, including mating flights, waggle dances, pupating larvae, flower pollination, and emerging queens. In some scenes, the high definition shots make already disturbing events even more disturbing, like bees dying after being exposed to chemicals and tiny varroa mites crawling around on the bodies of bees infecting them with diseases – wings wither away and bees become too weak to walk. This movie is worth watching for the impressive cinematography alone.

But bees aren’t the only actors. The human characters are almost as fun to watch. A Swiss beekeeper looks out over stunning views of the Alps where he keeps his bees. He follows a long tradition of beekeeping in his family and is very particular about maintaining a pure breed in his hives, going so far as flicking away the “wrong” bees from flowers on his property and crushing the head off of an unfaithful queen. A commercial beekeeper in the United States trucks thousands of beehives around the country, providing pollination services to a diverse group of farms – one of them being a massive almond grove in California. He has been witness to the loss of hundreds of honey bee colonies and has had to become “comfortable with death on an epic scale” – the grueling corporate world grinds along, and there is no time for mourning losses.

Further into the documentary, a woman in Austria demonstrates how she manipulates a colony into raising not just one queen, but dozens. She has spent years breeding bees, and her queens are prized throughout the world. A man in Arizona captures and raises killer bees – hybrid bees resulting from crosses between African and European honey bees (also known as Africanized honey bees). Despite their highly aggressive nature, he prefers them because they are prolific honey producers and they remain healthy without the use of synthetic pesticides.

Probably the darkest moment in the film is watching workers in China hand pollinate trees in an orchard. Excessive pesticide use has decimated pollinator populations in some regions, leaving humans to do the pollinating and prompting the narrator to reflect on the question, “Who’s better at pollinating, man or bees? Science answers with a definite, ‘not man.'”

Also included in the film is an intriguing discussion about bees as a super-organism with a German neuroscientist who is studying bee brains. The narrator sums it up like this: “Without its colony the individual bee cannot survive. It must subordinate its personal freedom for the good of the colony… Could it be that individual bees are like the organs or cells of a body? Is the super-organism as a whole the actual animal?”

Vanishing of the Bees

Colony collapse disorder is a sometimes veiled yet important theme throughout More than Honey, and it was certainly something that drove the creation of the film. In the case of Vanishing of the Bees, colony collapse disorder is the reason for its existence. Narrated by actor, Ellen Page, and produced in part by a film production company called Hive Mentality Films, this movie came out on the heels of the news that bee colonies were disappearing in record numbers throughout the world. It tells the story of colony collapse disorder from the time that it first appeared in the news – one of the film’s main characters is the beekeeper that purportedly first brought attention to the phenomenon – and into the years that followed as scientists began exploring potential causes.

This film contains lots of important information and much of it seems credible, but it is also the type of documentary that in general makes me wary of documentaries. Its purpose goes beyond just trying to inform and entertain; it’s also trying to get you on board with its cause. I may agree with much of what is being said, but I don’t particularly like having my emotions targeted in an effort to manipulate me to believe a certain way. It’s a good idea not to let documentaries or any other type of media form your opinions for you. Consider the claims, do some of your own research and investigation, and then come to your own conclusion. That’s my advice anyway…even though you didn’t ask for it.

That being said, colony collapse disorder is a serious concern, and so I’ll end by going back to More than Honey and leave you with this quote by its narrator:

The massive death of honey bees is no mystery. What’s killing them is not pesticides, mites, antibiotics, incest, or stress, but a combination of all these factors. They are dying as a result of our civilization’s success, as a result of man, who has turned feral bees into docile, domestic animals – wolves into delicate poodles.

March 14, 2015 by awkwardbotany

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.

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.

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:

How Stuff Works: How Bees Work – Included here are more excellent bee anatomy illustrations.
Understanding Bee Anatomy – This is a site I haven’t explored much, but it looks incredibly informative.
Bug Guide: Bees – Lots of bee photos and so much more.

Other “Year of Pollination” Posts:

January 14, 2015 by awkwardbotany

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)

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

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)

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)

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.

January 2, 2015 by awkwardbotany

2014: Year in Review

It is time again to look back at a year gone by and look forward to another year to come. Usually when we get to this point on the calendar, regardless of how my year has gone, I am anxious to put another year behind me and jump headlong into a new one, reinvigorated by that fresh start feeling that a new year seems to bring.

I manage this blog like a manage most things in my life, by the seat of my pants, not always sure where I am going with it but confident I will figure it out along the way. I have really enjoyed doing the blog this year, and I have felt a sense of direction for it emerging (at least in my mind; not sure if it comes across in the posts), and so in the spirit of that trajectory, I am thrilled to be entering my third year. I have a head full of ideas and I am gaining steam, so if things go the way I envision, this will be an abundant year of diverse posts that will hopefully prove to be enlightening, entertaining, and engrossing.

Serial Posts, etc.

In 2014 I started a few series of posts, and I plan to start more in 2015. The first one I started was an Ethnobotany series, which so far includes Holy Basil, Marigolds, and Cinchona. I also began a series on Drought Tolerant Plants, which so far consists of An Introduction, Fernbush, and Blue Sage. Flower Anatomy and Fruits were part of another new series exploring Botanical Terms. Some ideas for other series include: Poisonous Plants, Famous Botanists in History, and Botany in Popular Culture. None of these series has a regular posting schedule and each will continue indefinitely. I also plan to write more book reviews, as I only managed one in 2014 (Seedswap by Josie Jeffery). And speaking of reviews, probably my most ambitious endeavor of 2014 was reviewing the 17 articles in the October 2014 Special Issue of American Journal of Botany. You can read a recap here.

Social Media

It’s no mystery that having a social media presence in this day and age is imperative to the success of virtually any venture, especially a blog since the internet is veritably flooded with them. I’ve decided that Twitter is my favorite form of social media for now, and so I have been spending most of my time there. You can find and follow me @awkwardbotany. I also started a sister microblog on Tumblr in 2014. I mostly post plant and garden photos, and occasionally I share links to plant related things that I find interesting. Find and follow me here.

If you like what you read here and want to support Awkward Botany, the most helpful thing you can do is share it with your friends, family, and acquaintances. The easiest way to do that is by linking to individual posts on your preferred social media sites (there are buttons at the end of each post that help you do that). Or you can just tell people about it in person by using your mouth to make words, the old fashioned way. If you do share Awkward Botany online, consider including #phytocurious. You can also use this hashtag for anything plant related, including (especially) pictures of plants, that way I can easily find the cool plant things you are posting and share in your plant nerd glee.

Guest Posts

I hinted last year that I was considering publishing guest posts, and I have decided that I really want to do that. I’m going to be kind of picky about what I post, but don’t let that stop you from submitting something. You can write about your favorite plant, interesting plant science research, plants in the news, book or other media reviews, or anything else plant related. If this interests you, let me know by using the contact form or by sending me a message on Twitter. We can discuss further details from there.