Dung Moss (Revisited)

This is a revised version of a post that was originally published on January 14th, 2015. It includes excerpts from a chapter entitled, “Portrait of Splachnum,” in the book, Gathering Moss, by Robin Wall Kimmerer.

Certain plants, like corpse flowers and carrion flowers, emit foul odors when they bloom. The 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 employ stinky smells to recruit such insects to aid in reproduction and dissemination. Consider dung mosses.

No moss is more fastidious in its choice of habitats than Splachnum. Absent from the usual mossy haunts, Splachnum is found only in bogs. Not among the commoners like Sphagnum that build the peaty hummocks, not along the margins of the blackwater pools. Splachnum ampullaceum occurs in one, and only one, place in the bog. On deer droppings. On white-tailed deer droppings. On white-tailed deer droppings which have lain on the peat for four weeks. In July.

At least three genera (SplachnumTetraplodon, 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 strategy in which pollen or spores are distributed by insects. Compare this to anemophily, or wind pollination, which is the common way that moss spores are distributed. In fact, dung mosses are the only mosses known to exhibit entomophily.

Dung Moss (photo credit: wikimedia commons)

Dung Moss (photo credit: wikimedia commons)

Before we go too much further, it’s important to understand 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 plant parts. Lacking vascular tissues, water and nutrients are simply absorbed through the leaves and stems of mosses, which is why mosses are typically petite and prefer moist environments. Mosses also lack true roots and instead have rhizoids – threadlike structures that anchor the plants to their substrate of choice (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. In most plants, the haploid gametophytes are the sperm (pollen) and egg cells; the sporophyte is everything else. In mosses, the familiar green, leafy structure is actually the gametophyte. The gametophyte houses sperm and egg cells, and when the egg is fertilized by sperm it forms a zygote that develops into the sporophyte structure which extends above the leafy gametophyte. A capsule at the top of the sporophyte contains spores which are eventually released and, upon finding themselves on a suitable substrate in a hospitable environment, germinate to produce new plants. The spore then is comparable to a seed in vascular, seed-bearing plants.

photo credit: wikimedia commons

photo credit: wikimedia commons

As stated earlier, the spores of most mosses are distributed by wind. Dung mosses, on the other hand, employ flies in the distribution of their spores. They attract the flies by emitting scents that only flies can love from an area on the capsule of the sporophyte called the apophysis. This area is often enlarged and brightly colored in yellow, magenta, or red, giving it a flower-like appearance which acts as a visual attractant. The smells emitted vary depending on the type of substrate a particular species of dung moss inhabits. 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 rotting flesh, skeletal remains, or antlers.

Splachnum ampullaceum inhabits the droppings of white-tailed deer. Had a wolf or coyote followed the scent of the deer into the bog, its droppings would been colonized by S. luteum. The chemistry of carnivore dung is sufficiently distinct from that of herbivores to support a different species. … Moose droppings have their own loyal follower. The family to which Splachnum belongs includes several other mosses with an affinity for animal nitrogen. Tetraplodon and Tayloria can be found on humus, but primarily inhabit animal remains such as bones and owl pellets. I once found an elk skull lying beneath a stand of pines, with the jawbone tufted with Tetraplodon.

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

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

The set of circumstances that converge to bring Splachnum into the world is highly improbable. Ripening cranberries draw the doe to the bog. She stands and grazes with ears alert, flirting with the risk of coyotes. Minutes after she has paused, the droppings continue to steam. … The droppings send out an invitation written in wafting molecules of ammonia and butyric acid. Beetles and bees are oblivious to this signal, and go on about their work. But all over the bog, flies give up their meandering flights and antennae quiver in recognition. Flies cluster on the fresh droppings and lap up the salty fluids that are beginning to crystallize on the surface of the pellets. Gravid females probe the dung and insert glistening white eggs down into the warmth. Their bristles leave behind traces from their earlier foraging trips among the day’s dung, delivering spores of Splachnum on their footprints.

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 can 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 while receiving no nectar reward for their work.

There are 73 species in the Splachnaceae family, and nearly half of these species are dung mosses. Most are 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.

Since Splachnum can grow only on droppings, and nowhere else, the wind cannot be trusted with dispersal. Escape of the spores is successful only if they have both a means of travel and a reserved ticket for a particular destination. In the monotonous green of the bog, flies are attracted to the cotton candy colors of Splachnum, mistaking them for flowers. Rooting about in the moss for non-existent nectar the flies become coated with the sticky spores. When the scent of fresh deer droppings arrives on the breeze, the flies seek it out and leave Splachnum-coated footprints in the steaming dung.

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

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


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.

Grasshoppers – More Friend Than Foe?

Major outbreaks of grasshoppers can be devastating. A plague of locusts of biblical proportions can decimate crop fields and rangelands in short order. Clouds of grasshoppers moving in and devouring every plant in sight makes it easy to see why these insects are often seen as pests. Even small groups of them can do significant damage to a garden or farm. Yet, grasshoppers and their relatives have great ecological value and are important parts of healthy ecosystems. Love them or hate them, they are an essential piece of a bigger picture.

Grasshoppers are in the order Orthoptera, an order that includes katydids, crickets, wetas, and a few other familiar and not so familiar insects. Worldwide, there are more than 27,000 species of orthopterans. These insects mostly feed on plants; many are omnivorous while others are exclusively herbivorous. They are most commonly found in open, sunny, dry habitats like pastures, meadows, disturbed sites, open woods, prairies, and crop fields. Most insects in this order are fairly large, making them easy to identify; yet they don’t seem to receive the same level of human attention that charismatic insects like bees and butterflies do. In Field Guide to Grasshoppers, Katydids, and Crickets of the United States, the authors defend this diverse group of arthropods: “Grasshoppers often are thought of as modest-looking brown or green insects, but many species in this family are brightly colored, and some of the most dull-colored species rival butterflies in beauty when they spread their wings in flight.”

photo credit: wikimedia commons

photo credit: wikimedia commons

The voracious appetite of grasshoppers and their preference for plants can influence ecosystems in many ways. Certain plants may be favored over others, which affects the diversity and distribution of plant communities. Grasses are a particular favorite, despite being high in hard to digest compounds like lignin, cellulose, and silica. As grasshoppers consume vegetation – up to their body weight per day – digested materials return to the soil where soil dwelling organisms continue to break them down. In this way, grasshoppers and their relatives are major contributors to nutrient cycling. Returning nutrients to the soil results in increased nutrient availability for future plant growth. In fact, one grassland study found that despite short-term losses via grasshopper herbivory, plant growth was enhanced in the long-term due in part to accelerated nutrient cycling.

Because grasshoppers are such prolific consumers, their robust bodies are loaded with nutritious proteins and fats, making them a preferred food source for higher animals. Reptiles, raccoons, skunks, foxes, mice, and numerous species of birds regularly consume grasshoppers and related species. While many adult birds feed mostly on seeds and fruits, they seek out insects and worms to feed their young. Nutrient-packed grasshoppers are an excellent food source for developing birds. Humans in many parts of the world also find grasshoppers and crickets to be a tasty part of their diet.

Grasshoppers provide food for other invertebrates as well. The aforementioned field guide refers to the fate of grasshoppers and certain species of blister beetles as being “intimately linked,” because the larvae of these blister beetles feed exclusively on grasshopper eggs. Several species of flies and other insects, as well as spiders, also feed on grasshoppers and other orthopterans.

grasshopper on blade of grass

In short, grasshoppers play prominent roles in plant community composition, soil nutrient cycling, and the food chain. When grasshopper populations reach plague proportions, their impact is felt in other ways. From a human perspective, the damage is largely economic. However, their ability to thoroughly remove vegetation across large areas can be environmentally devastating as well, particularly when it comes to soil erosion and storm water runoff. The USDA’s Agricultural Research Service considers grasshoppers “among the most economically important pests” and cites research estimating that they are responsible for destroying as much as 23% of available range forage in the western United States annually. A paper published in the journal, Psyche, references a period between 2003-2005 in Africa where locusts were responsible for farmers losing as much as 80 to 100% of their crops.

This level of devastation is relatively rare. In Garden Insects of North America, Whitney Cranshaw states that of the more than 550 species of grasshoppers that occur in North America, “only a small number regularly damage gardens…almost all of these are in the genus Melanoplus.” Like most large, diverse groups of organisms, many grasshopper species are abundant and thriving while others are rare and threatened. Human activity has benefited certain species of grasshoppers while jeopardizing others. In general, grasshopper populations vary wildly from year to year depending on a slew of environmental factors.

Differential grasshopper (Melanoplus differentialis) - one of the four grasshoppers that Whitney Cranshaw lists as "particularly injurious" in his book Garden Insects of North America. (photo credit: www.eol.org)

Differential grasshopper (Melanoplus differentialis) – one of the four grasshoppers that Whitney Cranshaw lists as “particularly injurious” in his book Garden Insects of North America. (photo credit: www.eol.org)

A plague or outbreak of grasshoppers is a poorly understood phenomenon. It seems there are too many factors at play to pin such an occasion on any one thing. Warm, sunny, dry weather seems to favor grasshopper growth and reproduction, so drought conditions over a period of years can result in a dramatic increase in grasshopper populations. But drought can also limit plant growth, reducing the grasshoppers’ food supply. Natural enemies – which grasshoppers have many – also come into play. It seems that just the right conditions have to be met for an outbreak to occur – a seemingly unlikely scenario, but one that occurs frequently enough to cause concern.

Grasshoppers and fellow orthopterans are fascinating insects, and their place in the world is worth further consideration. For an example of just how compelling such insects can be, here is a story about crickets from Doug Tallamy’s book, Bringing Nature Home:

“Male tree crickets in the genus Oecanthus attempt to lure females to them by making chirping songs with their wings. The loudest male attracts the most females, so males often cheat a bit by positioning themselves within a cup-shaped leaf that amplifies the song beyond what the male can make without acoustical help. Each male chews a hole in the center of his cupped leaf that is just large enough to accommodate his raised wings during chirping. This ensures that the sound projects directly from the center of the parabolic leaf for maximum amplification.

Related Awkward Botany Posts:

The Making of a Kill Jar

I often hear stories from plant lovers about their initial nonchalance concerning plants. The common refrain seems to be that they were fascinated by wildlife and largely ignored plant life until they came to the realization that plants were integral in the lives of animals and play a major role in shaping the environments that support all life. Such an epiphany spawns an insatiable obsession with botany, at least for some people.

I seem to be on the opposite trajectory. It’s not like I have ever really been disinterested in animals; I’ve just been significantly more interested in plants and haven’t bothered to learn much about the animal kingdom (with the exception of entomology). My growing fascination with pollination biology (see last year’s Year of Pollination series) isn’t much of a stretch because insects have always appealed to me, and their intimate interactions with plants are hard to ignore. Ultimately, it is my interest in urban ecology and wildlife friendly gardening that is driving me to learn more about animals.

I started this year off by finally reading Doug Tallamy’s popular book, Bringing Nature Home. Tallamy wrote a lot about birds in his book, which got me thinking more about them. I then discovered Welcome to Subirdia, a book by John Marzluff that explores the diversity of birds that live among us in our urban environments. I then found myself paying more attention to birds. Many bird species rely on insects for food at some point in their lives. Plants regularly interact with insects both in defending themselves against herbivory and in attracting insects to assist in pollination. It’s all connected, and it seems I wouldn’t be much of a botanist then if I didn’t also learn about all of the players involved in these complex interactions.

So, now I’m a birdwatcher and an insect collector. Or at least I’m learning to be. Insects are hard to learn much about without capturing them. They often move quickly, making them hard to identify, or they go completely unnoticed because they are tiny and so well hidden or camouflaged. With the help of a net and a kill jar, you can get a closer look. This not only allows you to determine the species of insects that surround you, but it can also help give you an idea of their relative abundances, their life cycles, where they live and what they feed on, etc.

insect net 2_bw

As the name implies, if you’re using a kill jar, your actions will result in the death of insects. Some people will be more pleased about this than others. If killing insects bothers you, don’t worry, insect populations are typically abundant enough that a few individuals sacrificed for science will not hurt the population in a serious way.

Kill jars can be purchased or they can be made very simply with a few easy to find materials. Start with a glass jar with a metal lid. Mix up a small amount of plaster of paris. Pour the wet plaster in the jar, filling it to about one inch. Allow the plaster to dry completely. This process can be sped up by placing the jar in an oven set on warm. When the plaster is dry, “charge” the jar by soaking the plaster with either ethyl acetate, nail polish remover, or rubbing alcohol. I use nail polish remover because it is cheap and easily accessible. It doesn’t work as quickly as pure ethyl acetate, but it is less toxic. Place a paper towel or something soft and dry in the jar. This keeps the insects from getting beaten up too much as they thrash about. Once the insect is dead, it can be easily observed with a hand lens or a dissecting microscope. It can also be pinned, labeled, and added to a collection.

There are several resources online that describe the process of collecting and preserving insects, including instructions for making an inexpensive kill jar, which is why I am keeping this brief and will instead refer you to a couple of such sites. Like this one from Purdue University’s extension program. It’s directed toward youth, but it includes great information for beginners of any age. This post by Dragonfly Woman is a great tutorial for making a kill jar, and there are several other posts on her blog that are very informative for insect collectors of all experience levels.

I guess you could consider this part of my journey of becoming a naturalist. Perhaps you are on a similar journey. If so, share your thoughts and experiences in the comment section below.

Book Review: Bringing Nature Home

Since Bringing Nature Home by Douglas Tallamy was first published in 2007, it has quickly become somewhat of a “classic” to proponents of native plant gardening. As such a proponent, I figured I ought to put in my two cents. Full disclosure: this is less of a review and more of an outright endorsement. I’m fawning, really, and I’m not ashamed to admit it.


The subtitle pretty much sums it up: “How You Can Sustain Wildlife with Native Plants.” Ninety three pages into the book, Tallamy elaborates further: “By favoring native plants over aliens in the suburban landscape, gardeners can do much to sustain the biodiversity that has been one of this country’s richest assets.” And one of every country’s richest assets, as far as I’m concerned. “But isn’t that why we have nature preserves?” one might ask. “We can no longer rely on natural areas alone to provide food and shelter for biodiversity,” Tallamy asserts in the Q & A portion of his book. Humans have altered every landscape – urban, suburban, rural, and beyond – leaving species of all kinds threatened everywhere. This means that efforts to protect biodiversity must occur everywhere. This is where the You comes in. Each one of us can play a part, no matter how small. In closing, Tallamy claims, “We can each make a difference almost immediately by planting a native nearby.”

A plant is considered native to an area if it shares a historical evolutionary relationship with the other organisms in that area. This evolutionary relationship is important because the interactions among organisms that developed over thousands, even millions, of years are what define a natural community. Thus, as Tallamy argues, “a plant can only function as a true ‘native’ while it is interacting with the community that historically helped shape it.” A garden designed to benefit wildlife and preserve biodiversity is most effective when it includes a high percentage of native plants because other species native to the area are already adapted to using them.

Plants (and algae) are at the base of every food chain – the first trophic level – because they produce their own food using the sun’s energy. Organisms that are primarily herbivores are at the second trophic level, organisms that primarily consume herbivores are at the third trophic level, and so on. As plants have evolved they have developed numerous defenses to keep from being eaten. Herbivores that evolved along with those plants have evolved the ability to overcome those defenses. This is important because if herbivores can’t eat the plants then they can’t survive, and if they don’t survive then there will be little food for organisms at higher trophic levels.

The most important herbivores are insects simply because they are so abundant and diverse and, thus, are a major food source for species at higher trophic levels. The problem is that, as Tallamy learned, “most insect herbivores can only eat plants with which they share an evolutionary history.” Insects are specialized as to which plants they can eat because they have adapted ways to overcome the defenses that said plants have developed to keep things from eating them. Healthy, abundant, and diverse insect populations support biodiversity at higher trophic levels, but such insect populations won’t exist without a diverse community of native plants with which the insects share an evolutionary history.

That is essentially the thesis of Tallamy’s book. In a chapter entitled “Why Can’t Insects Eat Alien Plants?” Tallamy expounds on the specialized relationships between plants and insects that have developed over millennia. Plants introduced from other areas have not formed such relationships and are thus used to a much lesser degree than their native counterparts. Research concerning this topic was scarce at the time this book was published, but among other studies, Tallamy cites preliminary data from a study he carried out on his property. The study compared the insect herbivore biomass and diversity found on four common native plants vs. five common invasive plants. The native plants produced 4 times more herbivore biomass and supported 3.2 times as many herbivore species compared to the invasive plants. He also determined that the insects using the alien plants were generalists, and when he eliminated specialists from the study he still found that natives supported twice as much generalist biomass.

Apart from native plants and insects, Tallamy frames much of his argument around birds. Birds have been greatly impacted by humans. Their populations are shrinking at an alarming rate, and many species are threatened with extinction. Tallamy asserts, “We know most about the effects of habitat loss from studies of birds.” We have destroyed their homes and taken away their food and “filled their world with dangerous obstacles.” Efforts to improve habitat for birds will simultaneously improve habitat for other organisms. Most bird species rely on insects during reproduction in order to feed themselves and their young. Reducing insect populations by filling our landscapes largely with alien plant species threatens the survival of many bird species.

In the chapters “What Should I Plant?” and “What Does Bird Food Look Like?,” Tallamy first profiles 20 groups of native trees and shrubs that excel at supporting populations of native arthropods and then describes a whole host of arthropods and arthropod predators that birds love to eat. Tallamy’s fascinating descriptions of the insects, their life cycles, and their behaviors alone make this book worth reading. Other chapters that are well worth a look are “Who Cares about Biodiversity?” in which Tallamy explains why biodiversity is so essential for life on Earth, and “The Cost of Using Alien Ornamentals” in which Tallamy outlines a number of ways that our obsession with exotic plants has caused problems for us and for natural areas.

Pupa of ladybird beetle on white oak leaf (photo credit: wikimedia commons)

Pupa of a ladybird beetle on a white oak leaf. “The value of oaks for supporting both vertebrate and invertebrate wildlife cannot be overstated.” – Doug Tallamy (photo credit: wikimedia commons)

Convincing people to switch to using native plants isn’t always easy, especially if your argument involves providing habitat for larger and more diverse populations of insects. For those who are not fans of insects, Tallamy explains that “a mere 1%” of the 4 million insect species on Earth “interact with humans in negative ways.” The majority are not pests. It is also important to understand that even humans “need healthy insect populations to ensure our own survival.” Tallamy also offers some suggestions on how to design and manage an appealing garden using native plants. A more recent book Tallamy co-authored with fellow native plant gardening advocate Rick Darke called The Living Landscape expands on this theme, although neither book claims to be a how to guide.


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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 two

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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.