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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 (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 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)

References

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

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

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Year of Pollination: Pollinator Walk at Earthly Delights Farm

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

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

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

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

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

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

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

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

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

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

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

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

Bumblebee visiting Echinacea sp.

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

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

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

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Year of Pollination: Pollination Syndromes and Beyond

A discussion of pollination syndromes should begin with the caveat that they are a largely outdated way to categorize plant-pollinator interactions. Still, they are important to be aware of because they have informed so much of our understanding about pollination biology, and they continue to be an impetus for ongoing research. The concept of pollination syndromes exists in part because we are a pattern seeking species, endeavoring to place things in neat little boxes in order to make sense of them. This is relatively easy to do in a hypothetical or controlled environment where the parameters are selected and closely monitored and efforts are made to eliminate noise. However, the real world is considerably more dynamic than a controlled experiment and does not conform to black and white ways of thinking. Patterns are harder to unveil, and it takes great effort to ensure that observed patterns are genuine and not simply imposed by our pattern seeking brains.

That being said, what are pollination syndromes?  Pollination syndromes are sets of floral traits that are thought to attract specific types of pollinators. The floral traits are considered to have evolved in order to appeal to a particular group of pollinators – or in other words, selective pressures led to adaptations resulting in mutualistic relationships between plants and pollinators. Pollination syndromes are examples of convergent evolution because distantly related plant species have developed similar floral traits, presumably due to similar selection pressures. Pollination syndromes were first described by Italian botanist, Federico Delpino, in the last half of the 19th century. Over several decades his rudimentary ideas were fleshed out by other botanists, resulting in the method of categorization described (albeit briefly) below.

Honey bee on bee's friend (Phacelia tanacetifolia)

A honey bee getting friendly with bee’s friend (Phacelia tanacetifolia)

Pollination by bees (melittophily) – Flowers are blue, purple, yellow, or white and usually have nectar guides. Flowers are open and shallow with a landing platform. Some are non-symmetrical and tubular like pea flowers. Nectar is present, and flowers give off a mild (sometimes strong) sweet scent.

Pollination by butterflies (psychophily) – Flowers are pink, purple, red, blue, yellow, or white and often have nectar guides. They are typically large with a wide landing pad. Nectar is inside a long, narrow tube (or spur), and flowers have a sweet scent.

Pollination by hawkmoths and moths (sphingophily and phalaenophily) – Moth pollinated flowers open at night, have no nectar guides, and emit a strong, sweet scent. Flowers pollinated by hawkmoths are often white, cream, or dull violet and are large and tubular with lots of nectar. Those pollinated by other moths are smaller, not as nectar rich, and are white or pale shades of green, yellow, red, purple, or pink.

Pollination by flies (myophily or sapromyophily) – Flowers are shaped like a basin, saucer, or kettle and are brown, brown-red, purple, green, yellow, white, or blue.  Some have patterns of dots and stripes. If nectar is available, it is easily accessible. Their scent is usually putrid. A sapromyophile is an organism that is attracted to carcasses and dung. Flies that fall into this category visit flowers that are very foul smelling, offer no nectar reward, and essentially trick the fly into performing a pollination service.

Pollination by birds (ornithophily) –  Flowers are usually large, tubular, and red, orange, white, blue, or yellow. They are typically without nectar guides and are odorless since birds don’t respond to scent. Nectar is abundant and found at various depths within the flower.

Pollination by bats (chiropterophily) – Flowers are large, tubular or bell shaped, and white or cream colored with no nectar guides. They open at night, have abundant nectar and pollen, and have scents that vary from musty to fruity to foul.

Pollination by beetles (cantharophily) – Flowers are large and bowl shaped and green or white. There are no nectar guides and usually no nectar. The scent is strong and can be fruity, spicy, or putrid. Like flies, some beetles are sapromyophiles.

Locust borer meets rubber rabbitbrush (Ericameria nauseosa)

A locust borer meets rubber rabbitbrush (Ericameria nauseosa)

In addition to biotic pollination syndromes, there are two abiotic pollination syndromes:

Pollination by wind (anemophily) – Flowers are miniscule and brown or green. They produce abundant pollen but no nectar or odor. The pollen grains are very small, and the stigmas protrude from the flower in order to capture the windborne pollen.

Pollination by water (hydrophily) –  Most aquatic plants are insect-pollinated, but some have tiny flowers that release their pollen into the water, which is picked up by the stigmas of flowers in a similar manner to plants with windborne pollen.

This is, of course, a quick look at the major pollination syndromes. More complete descriptions can be found elsewhere, and they will differ slightly depending on the source. It’s probably obvious just by reading a brief overview that there is some overlap in the floral traits and that, for example, a flower being visited by a bee could also be visited by a butterfly or a bird. Such an observation explains, in part, why this method of categorizing plant-pollinator interactions has fallen out of favor. Studies have been demonstrating that this is not a reliable method of predicting which species of pollinators will pollinate certain flowers. A close observation of floral visitors also reveals insects that visit flowers to obtain nectar, pollen, and other items, but do not assist in pollination. These are called robbers. On the other hand, a plant species may receive some floral visitors that are considerably more effective and reliable pollinators than others. What is a plant to do?

Pollination syndromes imply specialization, however field observations reveal that specialization is quite rare, and that most flowering plants are generalists, employing all available pollinators in assisting them in their reproduction efforts. This is smart, considering that populations of pollinators fluctuate from year to year, so if a plant species is relying on a particular pollinator (or taxonomic group of pollinators) to aid in its reproduction, it may find itself out of luck. Considering that a flower may receive many types of visitors on even a semi-regular basis suggests that the selective pressures on floral traits may not solely include the most efficient pollinators, but could also include all other pollinating visitors and, yes, even robbers. This is an area where much more research is needed, and questions like this are a reason why pollination biology is a vibrant and robust field of research.

A bumble bee hugs Mojave sage (Salvia pachyphylla)

A bumble bee hugs the flower of a blue sage (Salvia pachyphylla)

Interactions between plants and pollinators is something that interests me greatly. Questions regarding specialization and generalization are an important part of these interactions. To help satiate my curiosity, I will be reading through a book put out a few years ago by the University of Chicago Press entitled, Plant-Pollinator Interactions: From Specialization to Generalization, edited by Nickolas M. Waser and Jeff Ollerton. You can expect future posts on this subject as I read through the book. To pique your interest, here is a short excerpt from Waser’s introductory chapter:

Much of pollination biology over the past few centuries logically focused on a single plant or pollinator species and its mutualistic partners, whereas a focus at the level of entire communities was uncommon. Recently we see a revival of community studies, encouraged largely by new tools borrowed from the theory of food webs that allow us to characterize and analyze the resulting patterns. For example, pollination networks show asymmetry – most specialist insects visit generalist plants, and most specialist plants are visited by generalist insects. This is a striking departure from the traditional implication of coevolved specialists!

References:

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Year of Pollination: Dung Moss

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

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

Dung Moss (photo credit: wikimedia commons)

Dung Moss (photo credit: wikimedia commons)

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

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

photo credit: wikimedia commons

photo credit: wikimedia commons

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

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

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

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

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

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

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

References

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

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